Influences on dusky dolphin (<Emphasis Type="Italic">Lagenorhynchus obscurus</Emphasis>) fission-fusion dynamics in Admiralty Bay,New Zealand

In societies characterized by a high degree of fission-fusion dynamics, individuals adjust their grouping patterns according to the shifting balance of costs and benefits associated with grouping. This study examines influences on fission-fusion dynamics for dusky dolphins (Lagenorhynchus obscurus) in Admiralty Bay, New Zealand. This area is an important foraging habitat for dusky dolphins during the winter and spring. Admiralty Bay has little predation risk, but nearshore mussel farms may infringe on available habitat. I used generalized estimating equations to determine the influences of coordinated foraging, predation risk, and presence of mussel farms on party size, rate of fission-fusion, and behavioral state. I conducted 168 boat-based group focal follows totaling 168 h. The proportion of individuals observed foraging was positively related to party size and rate of party fusion. Resting had no effect on party size and did not vary according to location. Near mussel farms, traveling decreased, and rate of party fission decreased. I conclude that (1) coordinated foraging strategies are a primary influence on fission-fusion dynamics within this population, (2) dolphins may respond to decreased predation risk by not adjusting party size or location during resting, and (3) areas near mussel farms are not used for traveling.     

Fissioning minimizes ranging costs in spider monkeys: a multiple-level approach

The adjustment to deal with intragroup food competition is probably the most plausible explanation of high levels of fission–fusion dynamics. However, studies did not always support expected relations between food availability, ranging costs, and subgroup size. We used several levels of analysis differing in the time and spatial scale in order to investigate this explanation in spider monkeys. In our study, subgroups were larger when food availability was higher across most levels of analyses used. We also found a fine-scale adjustment: compared to the food patch previously visited, spider monkeys traveled to larger patches just after fusions. This was not without an immediate travel cost: the interpatch distance and travel time after a fusion were longer than that before the fusion. This rapid adjustment shows the flexibility that fission–fusion dynamics can offer. Spider monkeys are in large subgroups only when food conditions are favorable, as evidenced by the fact that at all the other time-scale levels larger subgroups did not experience greater ranging costs than smaller subgroups. Our results indicate that on the whole spider monkeys successfully minimize ranging costs by fission and fusion of subgroups.     

I know you: familiarity with an audience influences male-male interactions in Siamese fighting fish, Betta splendens

The presence of an audience alters male–male interactions in a number of social species. Additionally, whether individuals have interacted previously may influence the current encounter. Although it is clear that previous exposure influences communication and that the presence of an audience affects behavior in a number of contexts, how these phenomena might interact to impact interactions is underexplored. This study examined the effects of audience familiarity on male–male interactions in Siamese fighting fish (Betta splendens). Treatments that varied based on familiarity with and sex of the audience as well as length of time after initial audience exposure were used to investigate whether male aggressive interactions were affected by these factors. Males first encountered an individual that would later become an audience. Pairs of males then interacted with each other in either the presence of this familiar individual or a new, unfamiliar audience conspecific or with no audience at all. Opponent-directed gill flaring and tail beats were influenced by familiarity with the audience individual, although they were affected differently by the sex of the audience present. Gill flaring was affected predominantly by the presence of a male audience, whereas tail beat rate was influenced by the presence of a female audience. Gill flaring rate did not differ regardless of whether 4 or 24?h had passed between the initial exposure to the audience and the testing phase. In contrast, the familiarity effect disappeared after 4?h for tail beats. These results stress the importance of considering the familiarity of individuals within a network when examining communication networks.     

Growth dynamics and mortality of the intertidal encrusting sponge Halichondria okadai (Demospongiae, Halichondrida)

A periodical survey of the growth of an intertidal encrusting sponge Halichondria okadai was conducted on a rocky intertidal shore on the Izu Peninsula, southern Japan. The area covered by each of 16 marked sponges was monitored, and the fusion and fission of specimens were recorded from June 1995 to June 1998. During the investigation, 15 of the original specimens managed to survive accompanying fusions or fissions and only one colony completely disappeared. Ten fusions and 23 fissions were observed in total. The maximum death rate during a semilunar period was 16.7% and cumulative mortality throughout the study period was 46.1%, indicating that the mortality of H. okadai is lower than that of other species previously reported. There was no particular seasonal pattern in the occurrences of fusion, fission and death of sponges. However, fissions were frequently observed in the latter half of the study period and were sometimes followed by fusions or death of specimens. Positive growth rates were often observed during warmer months, and specimens showed little growth or regression in winter. Seasonal patterns in surface area were thought to be related to seasonality in reproductive activity.     

To fission or to fusion: effects of community size on wild chimpanzee (<Emphasis Type="Italic">Pan troglodytes verus</Emphasis>) social organisation

To balance advantages and disadvantages of group living, some species have fission-fusion social systems in which members of the same group form frequently changing subgroups. This allows flexible responses of group size to external conditions while at the same time retaining group stability. In chimpanzees, subgroup (party) size and composition depend mainly on the presence of receptive females, food availability and the activity of the party. Here we analyse the extent to which fission-fusion parameters are influenced by changes of demographic variables like community size and composition. Data were collected from a habituated West African chimpanzee community (Taï forest, Côte dIvoire) over 10 years, during which total community size decreased from 51 to 21, and the number of adult males decreased from 9 to 2. Taï chimpanzees are highly gregarious, as they spend more than 80% of their time with unrelated conspecifics. With decreasing community size, party size, party duration and male-female association increased. Neither activity nor the presence of receptive females or feeding competition could explain the observed changes in grouping patterns. Thus, the decrease in community size led to an increase of party cohesion and also enhanced cohesiveness between the sexes, while general sociality remained unchanged. Therefore, our data support the notion that small communities are more cohesive and have a less flexible fission-fusion system.Communicated by D. Watts     

Offspring sex ratios reflect lack of repayment by auxiliary males in a cooperatively breeding passerine

The repayment hypothesis posits that primary sex ratios in cooperative species should be biased towards the helping sex because these offspring “repay” a portion of their cost through helping behavior and therefore are less expensive to produce. However, many cooperatively breeding birds and mammals do not show the predicted bias in the primary sex ratio. Recent theoretical work has suggested that the repayment hypothesis should only hold when females gain a large fitness advantage from the presence of auxiliary adults in the group. When auxiliaries provide little or no fitness advantage, competition between relatives should lead to sex ratios biased towards the dispersing (non-helping) sex. We examined the benefits auxiliaries provide to females and corresponding offspring sex ratios in the red-backed fairy-wren (Malurus melanocephalus), a cooperatively breeding Australian bird with male auxiliary helpers. We found that auxiliaries provide little or no benefit to female reproductive success or survival. As predicted, the population primary sex ratio was biased towards daughters, the dispersing sex, and females with auxiliaries produced female-biased broods whereas females without auxiliaries produced unbiased broods. Moreover, offspring sex ratios were more strongly biased toward females in years when auxiliaries were more common in the population. These results suggest that offspring sex ratios are associated with competition among the non-dispersing sex in this species, and also that females may use cues to assess local breeding opportunities for their offspring.     

Ecological constraints on group size: an analysis of spider monkey and chimpanzee subgroups

The social organization of spider monkeys (Ateles geoffroyi) and chimpanzees (Pan troglodytes) appear remarkably similar. In this paper, field studies of these two species were used to (1) test a model of ecological constraints on animal group size which suggests that group size is a function of travel costs and (2) assess ecological and social factors underlying the social organization of these two species. Spider monkeys were studied over a 6-year period in Santa Rosa National Park, Costa Rica, and chimpanzees were studied for 6 years in Kibale National Park, Uganda. Adults of both species spent their time in small subgroups that frequently changed size and composition. Thus, unlike most primate species, spider monkeys and chimpanzees were not always in a spatially cohesive social group; each individual had the option of associating in subgroups of a different size or composition. Both species relied on ripe fruit from trees that could be depleted through their feeding activity. However, spider monkey food resources tended to occur at higher densities, were more common, less temporally variable, and did not reach the low levels experienced by chimpanzees. Analyses of the relationship between subgroup size and the density and distribution of their food resources suggested that travel costs limit subgroup size. However, these ecological factors did not influence all age/sex classes equally. For example, the number of adult males in a subgroup was a function of food density and travel costs. However, this was not the case for female chimpanzees, suggesting that the benefits of being in a subgroup for females did not exceed the costs, even when ecological conditions appeared to minimize subgroup foraging costs. Therefore, it seems likely that social strategies influenced the relationship between food resource variables and subgroup size.     

Loud calls as a mechanism of social coordination in a fission–fusion taxon, the white-bellied spider monkey (Ateles belzebuth)

Spider monkeys (Ateles spp.) live in social groups that exhibit high levels of fission–fusion dynamics, in which group members form subgroups of varying sizes and compositions. Within these fluid societies, how individuals establish contact with dispersed group members with whom they might choose to associate remains unclear. Long-range vocalizations might facilitate interactions between group members and provide a means of social coordination in fission–fusion societies. We evaluated this possibility for one spider monkey vocalization, the loud call, by examining calling behavior, the relationship between loud calls and changes in subgroup size, and the response of individuals to distant calls and playback experiments in a single study group. We found that 82 % of loud calls were emitted within 30 min of a call from a different location, suggesting that individuals frequently emit loud calls in response to the calls of distant group members. Subgroups that emitted loud calls, especially those that responded to distant calls, were much more likely to experience an increase in subgroup size within an hour after calling than those that did not. Animals also approached distant loud calls more than they avoided or ignored these calls. Finally, playbacks of male calls demonstrated that females respond preferentially to the calls of some individuals over others. Taken together, these results provide support for the hypothesis that spider monkey loud calls function to facilitate and initiate interactions between dispersed group members and suggest that vocal signals can play an important role in influencing social interactions in fission–fusion societies.     

Optimal movement strategies for social foragers in unpredictable environments

Spatial movement models often base movement decision rules on traditional optimal foraging theories, including ideal free distribution (IFD) theory, more recently generalized as density-dependent habitat selection (DDHS) theory, and the marginal value theorem (MVT). Thus optimal patch departure times are predicted on the basis of the density-dependent resource level in the patch. Recently, alternatives to density as a habitat selection criterion, such as individual knowledge of the resource distribution, conspecific attraction, and site fidelity, have been recognized as important influences on movement behavior in environments with an uncertain resource distribution. For foraging processes incorporating these influences, it is not clear whether simple optimal foraging theories provide a reasonable approximation to animal behavior or whether they may be misleading. This study compares patch departure strategies predicted by DDHS theory and the MVT with evolutionarily optimal patch departure strategies for a wide range of foraging scenarios. The level of accuracy with which individuals can navigate toward local food sources is varied, and individual tendency for conspecific attraction or repulsion is optimized over a continuous spectrum. We find that DDHS theory and the MVT accurately predict the evolutionarily optimal patch departure strategy for foragers with high navigational accuracy for a wide range of resource distributions. As navigational accuracy is reduced, the patch departure strategy cannot be accurately predicted by these theories for environments with a heterogeneous resource distribution. In these situations, social forces improve foraging success and have a strong influence on optimal patch departure strategies, causing individuals to stay longer in patches than the optimal foraging theories predict.     

Genetic relationships between roost-mates in a fission–fusion society of tree-roosting big brown bats (<Emphasis Type="Italic">Eptesicus fuscus</Emphasis>)

Kin-based patterns of associations are often observed in group living mammals. Colonies of forest-living big brown bats (Eptesicus fuscus) exhibit fission–fusion roosting behavior and female philopatry. Within a roosting area of forest, adult females are distributed into several subgroups roosting in different trees during the day. At night, adult females leave the roost subgroups to forage and, upon return to the roosting area at dawn, both the individual composition and location of subgroups often change. Individuals exhibit nonrandom roosting associations, and we hypothesized that genetic relationships would influence roosting associations. We determined (1) whether the strength of roosting associations between pairs of bats (based on radiotelemetry) was correlated with relatedness, (2) whether individuals that roosted together in roost subgroups were more related than by chance, and (3) from roost subgroups, the pairs of bats that roosted nonrandomly and whether the proportion of related pairs was higher than expected at random. Relatedness measures were based on microsatellite genotyping and mitochondrial DNA sequences. We found from all analyses that roosting associations were not influenced by relatedness or matrilineal relationships. These results provide clear evidence that, contrary to other mammals, kinship does not mediate roosting associations within forest living big brown bats that exhibit fission–fusion roosting behavior.     

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.     

Males are sensitive — sex-dependent effect of rearing conditions on nestling growth

The sex-dependent effect of environmental conditions on nestlings has been extensively studied in size dimorphic birds. Whether males or females are more sensitive to poor conditions is not yet clear; however, the degree of sexual size-dimorphism, brood size and their interactions seem to influence the pattern. Much less is known about sex-dependent environmental sensitivity in size-monomorphic species, even though it may result in biased sex allocation. We altered the rearing conditions by brood size manipulation in the size-monomorphic collared flycatcher and then examined the sex-specific development of the nestlings. In all analyses, we controlled for the effect of paternity, because one may expect extra-pair young to be of better genetic quality and perform better at least under poor conditions. However, this was not the case, because we did not find any difference in growth rate or fledging size between extra- and within-pair young. We found that male nestlings had the potential for faster growth under favourable conditions, but suffered more under poor conditions. We found no sex × environment interaction for fledging size probably because the growth curves level off before fledging, and the disadvantaged nestlings can catch up with their siblings. The larger sensitivity of males does not explain the previously found seasonal shift in brood sex ratios and contradicts previous findings in another size-monomorphic species where females were more sensitive. This suggests that even in size-monomorphic species, no general rule exists, which determines the more sensitive sex.     

Wildlife contact analysis: emerging methods, questions, and challenges

Recent technological advances, such as proximity loggers, allow researchers to collect complete interaction histories, day and night, among sampled individuals over several months to years. Social network analyses are an obvious approach to analyzing interaction data because of their flexibility for fitting many different social structures as well as the ability to assess both direct contacts and indirect associations via intermediaries. For many network properties, however, it is not clear whether estimates based upon a sample of the network are reflective of the entire network. In wildlife applications, networks may be poorly sampled and boundary effects will be common. We present an alternative approach that utilizes a hierarchical modeling framework to assess the individual, dyadic, and environmental factors contributing to variation in the interaction rates and allows us to estimate the underlying process variation in each. In a disease control context, this approach will allow managers to focus efforts on those types of individuals and environments that contribute the most toward super-spreading events. We account for the sampling distribution of proximity loggers and the non-independence of contacts among groups by only using contact data within a group during days when the group membership of proximity loggers was known. This allows us to separate the two mechanisms responsible for a pair not contacting one another: they were not in the same group or they were in the same group but did not come within the specified contact distance. We illustrate our approach with an example dataset of female elk from northwestern Wyoming and conclude with a number of important future research directions.     

Divided destinies: group choice by female savannah baboons during social group fission

Group living provides benefits to individuals while imposing costs on them. In species that live in permanent social groups, group division provides the only opportunity for nondispersing individuals to change their group membership and improve their benefit to cost ratio. We examined group choice by 81 adult female savannah baboons (Papio cynocephalus) during four fission events. We measured how each female’s group choice was affected by several factors: the presence of her maternal kin, paternal kin, age peers, and close social partners, her average kinship to groupmates, and her potential for improved dominance rank. Maternal kin, paternal kin, and close social partners influenced group choice by some females, but the relative importance of these factors varied across fissions. Age peers other than paternal kin had no effect on group choice, and average kinship to all groupmates had the same effect on group choice as did maternal kin alone. Most females were subordinate to fewer females after fissions than before, but status improvement did not drive female group choice; females often preferred to remain with social superiors who were their close maternal kin, rather than improving their own social ranks. We suggest that during permanent group fissions, female baboons prefer to remain with close maternal kin if those are abundant enough to influence their fitness; if they have too few close maternal kin then females prefer to remain with close paternal kin, and social bonds with nonkin might also become influential. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Social network analysis resolves temporal dynamics of male dominance relationships

Social organization is often studied through point estimates of individual association or interaction patterns, which does not account for temporal changes in the course of familiarization processes and the establishment of social dominance. Here, we present new insights on short-term temporal dynamics in social organization of mixed-sex groups that have the potential to affect sexual selection patterns. Using the live-bearing Atlantic molly (Poecilia mexicana), a species with pronounced male size polymorphism, we investigated social network dynamics of mixed sex experimental groups consisting of eight females and three different-sized males over a period of 5 days. Analyzing association-based social networks as well as direct measures of spatial proximity, we found that large males tended to monopolize most females, while excluding small- and medium-bodied males from access to females. This effect, however, emerged only gradually over time, and different-sized males had equal access to females on day 1 as well as day 2, though to a lesser extent. In this highly aggressive species with strong social dominance stratifications, the observed temporal dynamics in male-female association patterns may balance the presumed reproductive skew among differentially competitive male phenotypes when social structures are unstable (i.e., when individual turnover rates are moderate to high). Ultimately, our results point toward context-dependent sexual selection arising from temporal shifts in social organization.     

Parentally biased favouritism in relation to offspring sex in zebra finches

Family conflicts over parental care result in offspring attempting to exert control using solicitation behaviours, whilst the parents are potentially able to retaliate through provisioning rules. However, the evolutionary interests of one parent may not necessarily support the evolutionary interests of the other parent, and such conflicts of interest may be expressed in how the two parents allocate the same form of parental care to individual offspring. Theory suggests that such parentally biased favouritism is a universally predicted outcome of evolutionary conflicts of interest, and empirical evidence suggests that parentally biased favouritism occurs in relation to offspring size and solicitation behaviours. However, unequivocal empirical evidence of parentally biased favouritism in relation to offspring sex is absent, due to being strongly confounded by sex differences in size and solicitation behaviours. Here, we present strong evidence for parentally biased favouritism in relation to offspring sex in zebra finches (Taeniopygia guttata), independent of the effects of chick size and begging intensity. Mothers preferentially provisioned sons over daughters, whilst fathers showed no bias, meaning that sons received more food than daughters. Parentally biased favouritism in relation to offspring sex facilitates parental control over evolutionary conflicts of interest and is probably more widespread than previously realised.     

Spatial scale of local breeding habitat quality and adjustment of breeding decisions

Experimental studies provide evidence that, in spatially and temporally heterogeneous environments, individuals track variation in breeding habitat quality to adjust breeding decisions to local conditions. However, most experiments consider environmental variation at one spatial scale only, while the ability to detect the influence of a factor depends on the scale of analysis. We show that different breeding decisions by adults are based on information about habitat quality at different spatial scales. We manipulated (increased or decreased) local breeding habitat quality through food availability and parasite prevalence at a small (territory) and a large (patch) scale simultaneously in a wild population of Great Tits (Parus major). Females laid earlier in high-quality large-scale patches, but laying date did not depend on small-scale territory quality. Conversely, offspring sex ratio was higher (i.e., biased toward males) in high-quality, small-scale territories but did not depend on large-scale patch quality. Clutch size and territory occupancy probability did not depend on our experimental manipulation of habitat quality, but territories located at the edge of patches were more likely to be occupied than central territories. These results suggest that integrating different decisions taken by breeders according to environmental variation at different spatial scales is required to understand patterns of breeding strategy adjustment.     

Population structure and sex-change in the coral-inhabiting snailCoralliophila violacea at Hsiao-Liuchiu,Taiwan

Surveys of the coral-inhabiting snailCoralliophila violacea (Lamarck) (=C. neritoidea Kiener) were made on shallow fringing reefs (<8 m deep) around Hsiao-Liuchiu, Taiwan, between July and October 1990. The snails were aggregated into patches on the surface of massive poritid coral colonies. Coral colonies >40 cm in diameter were more likely to bear patches of snails than smaller colonies, and also to have more snails. The coralliophilids ranged from 5 to 30 mm in aperture length. The sex ratio of the population was biased toward males (539:279), with only a few small individuals of indistinguishable sex. Snails between 6 and 10 mm were all males, while most snails with aperture lengths 20 mm were females. Judging from the distinct size ranges of males and females within patches and from the observed degeneration of the penis, the snails may have changed sex from male to female with increasing size. Sex-change may occur across a wide size range (10 to 20 mm). The correlation of smallest female size and largest male size among patches indicates that snail size at sex-change is peculiar to each individual patch. Those females in patches with a single female (but many males) were significantly smaller than females in multiple-female patches. It is likely that in the absence of females males change sex at a smaller size, whereas in the presence of large females males delay sexchange until they have reached a larger size. The plasticity of size at sex-change may be adaptive and a result of natural selection at the individual level.