Fission–Fusion Dynamics in Southern Muriquis (Brachyteles arachnoides) in Continuous Brazilian Atlantic Forest

Fission–fusion social dynamics, the spatiotemporal variance in subunit size, composition, and cohesion, are the subject of considerable interest owing to their complex nature and widespread appearance in the primate order. We here aim to describe the nature of fission–fusion dynamics in a population of egalitarian southern muriquis inhabiting a relatively undisturbed extension of mildly seasonal Brazilian Atlantic forest to provide insights into the functions of fission–fusion dynamics and to examine the constraints on and opportunities for associations among individuals. We collected instantaneous scan samples and ad libitum data over a total of 13 mo in 2 yr (1599 observation hours) on subunit size at 2 spatiotemporal scales (party and nomadic party), party composition, spatial cohesion, and the behavioral context of fission–fusion events. These southern muriquis exhibited high levels of fission–fusion with significant variation in party size, cohesion, and composition. The group was weakly cohesive with a small mean party size (3.74 adults, 5.32 all individuals) and nomadic party size (13.73 adults, 19.38 all individuals). Mixed parties were the most frequently observed party type, although the high frequency of all-male parties suggests strong relationships based on philopatry and a mating strategy under scramble competition. We compare fission–fusion dynamics across populations of Brachyteles and highlight the striking continuum of these dynamics in the genus. We make interspecific comparisons with Ateles and Pan, with well documented high levels of fission–fusion, and demonstrate interspecific variation and convergence in grouping patterns.     

Dusky dolphin group dynamics and association patterns in Península Valdés,Argentina

We inspected dusky dolphin (Lagenorhynchus obscurus) group dynamics in Golfo Nuevo and found differences in social organization between cold and warm seasons. Surveys were conducted onboard a research vessel, from which we collected behavioral observations and group fission‐fusion data from 2001 to 2008; we also collected photo‐identification data from 2004 to 2012. To analyze association patterns, we calculated half‐weight association index (HWI) and social differentiation (S). We conducted a Monte Carlo permutation test to determine whether observed association patterns were significantly different from random association using a compiled version of SOCPROG 2.7. Group fission and fusion dynamics depended on group behavior, the main activity after the groups' fusion was feeding, and they never fission before socializing. The social structure of dusky dolphins included long‐term preferred companions in the cold season; during the warm season, there were no preferred companions. This seasonal difference in social structure could be related to an accompanying shift in foraging behaviors that appears to be driven by changes in prey availability. If so, then a loosening of bonds among individuals during the warm season, when prey is more available, would reflect these social structure changes.     

Dusky dolphin (Lagenorhynchus obscurus) social structure characterized by social fluidity and preferred companions

Many coastal delphinids exhibit social structures with a high degree of fission‐fusion dynamics. However, little is known about fission‐fusion dynamics in semipelagic delphinids such as dusky dolphins (Lagenorhynchus obscurus). We analyzed dusky dolphin social structure in a winter foraging habitat in New Zealand. We examined the population‐level societal attributes of preferred/avoided companions, behaviorally specific preferred/avoided companions, and simple ratio (SR) association index strength and distribution and the dyad‐level societal attribute of temporal patterning of associations. We identified 741 individuals; 99 met inclusion criteria for analysis. Long‐term and short‐term preferred/avoided companions were present in the pooled (6 yr) and annual samples. Individuals were observed to form long‐term but not short‐term preferred/avoided companions during all behavioral states. Mean and maximum SRs were 0.03 ± 0.01 SD and 0.40 ± 0.20 SD, respectively. Temporal analysis indicated that individuals associated preferentially throughout the study period. Dusky dolphins exhibited high social fluidity, as indicated by many weak and few strong associations. However, there was stability as individuals were observed to form behaviorally specific preferred/avoided companions and associate preferentially throughout the 6 yr period. Dusky dolphins thus display a dynamic social structure where overall low levels of bonding are juxtaposed against a subset of the population that forms strong bonds.     

Group composition influences on behavioral sequence patterns of the Guiana dolphin Sotalia guianensis

Analysis of behavioral sequences displayed by different individuals can be used to infer individual decisions under diverse conditions. This study investigates whether behavioral transitions and routines differ according to the group composition of Guiana dolphins (Sotalia guianensis). We used the Z-score binomial test to detect significant behavioral transitions from a first-order transition matrix. The directed tree method was used to identify behavioral routines. Solitary individuals presented fewer transitions than heterogeneous groups. More routines were observed for heterogeneous than for homogeneous groups, and fewer routines for solitary individuals. Homogeneous groups maximized routines ending in traveling and milling, while heterogeneous groups presented higher probabilities of socializing. The analyses indicate that subgroups formed during Guiana dolphin fission–fusion events differed in probability of behavioral sequences, reflecting different feeding, resting, and socializing needs at different moments.     

Seasonal and sex differences in the fission–fusion dynamics of spider monkeys (<Emphasis Type="Italic">Ateles geoffroyi yucatanensis</Emphasis>) in Belize

Spider monkeys (Ateles sp.) are characterized by high fission–fusion dynamics, meaning their social grouping pattern is fluid and consists of subgroups that vary in size, composition, and spatial cohesion over time. In this study, we quantify the fission–fusion dynamics of a group of spider monkeys at Runaway Creek Nature Reserve in Belize by measuring subgroup size, spatial cohesion, and stability using data spanning 5 years. We then test whether variation in these three subgroup measures differ according to season, subgroup sex composition, and the reproductive status of female subgroup members. Our results show that subgroups were larger in size and less stable in membership during the wet season compared to the dry season. All-female subgroups were less spatially cohesive but more stable in membership than all-male subgroups. Finally, we report that subgroups with one or more non-lactating females (i.e., without nursing young) were smaller on average than subgroups containing lactating females with nursing young. These data contribute to a growing body of research documenting the ecological and social dimensions along which grouping patterns might vary.     

Whistle variation in Mediterranean common bottlenose dolphin: The role of geographical,anthropogenic, social,and behavioral factors

The studies on the variation of acoustic communication in different species have provided insight that genetics, geographic isolation, and adaptation to ecological and social conditions play important roles in the variability of acoustic signals. The dolphin whistles are communication signals that can vary significantly among and within populations. Although it is known that they are influenced by different environmental and social variables, the factors influencing the variation between populations have received scant attention. In the present study, we investigated the factors associated with the acoustic variability in the whistles of common bottlenose dolphin (Tursiops truncatus), inhabiting two Mediterranean areas (Sardinia and Croatia). We explored which factors, among (a) geographical isolation of populations, (b) different environments in terms of noise and boat presence, and (c) social factors (including group size, behavior, and presence of calves), were associated with whistle characteristics. We first applied a principal component analysis to reduce the number of collinear whistle frequency and temporal characteristics and then generalized linear mixed models on the first two principal components. The study revealed that both geographic distance/isolation and local environment are associated with whistle variations between localities. The prominent differences in the acoustic environments between the two areas, which contributed to the acoustic variability in the first principal component (PC1), were found. The calf's presence and foraging and social behavior were also found to be associated with dolphin whistle variation. The second principal component (PC2) was associated only with locality and group size, showing that longer and more complex tonal sound may facilitate individual recognition and cohesion in social groups. Thus, both social and behavioral context influenced significantly the structure of whistles, and they should be considered when investigating acoustic variability among distant dolphin populations to avoid confounding factors.     

Social and spatial effects on genetic variation between foraging flocks in a wild bird population

Social interactions are rarely random. In some instances, animals exhibit homophily or heterophily, the tendency to interact with similar or dissimilar conspecifics, respectively. Genetic homophily and heterophily influence the evolutionary dynamics of populations, because they potentially affect sexual and social selection. Here, we investigate the link between social interactions and allele frequencies in foraging flocks of great tits (Parus major) over three consecutive years. We constructed co‐occurrence networks which explicitly described the splitting and merging of 85,602 flocks through time (fission–fusion dynamics), at 60 feeding sites. Of the 1,711 birds in those flocks, we genotyped 962 individuals at 4,701 autosomal single nucleotide polymorphisms (SNPs). By combining genomewide genotyping with repeated field observations of the same individuals, we were able to investigate links between social structure and allele frequencies at a much finer scale than was previously possible. We explicitly accounted for potential spatial effects underlying genetic structure at the population level. We modelled social structure and spatial configuration of great tit fission–fusion dynamics with eigenvector maps. Variance partitioning revealed that allele frequencies were strongly affected by group fidelity (explaining 27%–45% of variance) as individuals tended to maintain associations with the same conspecifics. These conspecifics were genetically more dissimilar than expected, shown by genomewide heterophily for pure social (i.e., space‐independent) grouping preferences. Genomewide homophily was linked to spatial configuration, indicating spatial segregation of genotypes. We did not find evidence for homophily or heterophily for putative socially relevant candidate genes or any other SNP markers. Together, these results demonstrate the importance of distinguishing social and spatial processes in determining population structure.     

Social communities and spatiotemporal dynamics of association patterns in estuarine bottlenose dolphins

Network analysis has recently been used to delve into the dynamics of cetacean sociality. Few studies, however, have addressed how habitat shape influences sociality, specifically how linear water bodies constrain the space where individuals might interact. We utilized network and spatiotemporal analyses to investigate association patterns and community structure in a bottlenose dolphin population in a linear estuarine system, the Indian River Lagoon (IRL), Florida. Using sighting histories from a multiyear photo‐identification study we examined association patterns for 185 individuals collected over a 6.5 yr period (2002–2008). The population was highly differentiated (S = 0.723) and organized into six distinct social communities (Q = 0.544), spread in an overlapping pattern along the linear system. Social organization differed between communities, with some displaying highly interconnected networks and others comprising loosely affiliated individuals with more ephemeral associations. Temporal patterns indicated short‐term associations were a significant feature of the fission‐fusion dynamics of this population. Spatial analyses revealed that social structure was shaped by an individual's ranging patterns and by social processes including preference and avoidance behavior. Finally, we found that habitat “narrowness” may be a major driving force behind the sociality observed.     

The Dynamic of Aggression: How Individual and Group Factors Affect the Long‐Term Interspecific Aggression Between Two Sympatric Species of Dolphin

Interspecific aggression, similar to intergroup conspecific aggression, has been observed in a variety of taxa. The dominant group or individual is determined by multiple aggressive events and can be influenced by the size, age, or group size of the participating individuals. Interspecific aggression between Atlantic bottlenose (Tursiops truncatus) and spotted (Stenella frontalis) dolphins, both resident and sympatric to Little Bahama Bank, the Bahamas has been consistently observed for over two decades. However, it is unclear whether one species is more dominant and little is known about the factors that influence the progression of aggression. For this study, underwater video recordings of 32 aggressive encounters composed of 451 aggressive behavioural events were analysed over a 12‐yr period (1993–2004). These were used to describe the interspecific aggression observed and quantify which factors (the species and age class of the participants or the group size and behaviour of spotted dolphin groups) had the strongest impact on the progression and outcome of aggression. Over the long term, interspecific aggression was bidirectional with neither species being more dominant. During a single encounter, spotted dolphin group synchrony had the strongest impact on the dynamic of aggression, specifically impacting which group (1) initiated aggression, (2) the direction of aggression and (3) the occurrence of dynamic shifts or dominance reversals. This is the first study to quantify the dynamic of aggression for this population, to document bidirectional aggression and dynamic shifts during long‐term interspecific aggression in free‐ranging delphinids, and this study quantifies the role of synchrony during interspecific aggression using underwater observations.     

To follow or not? How animals in fusion–fission societies handle conflicting information during group decision‐making

When group members possess differing information about the environment, they may disagree on the best movement decision. Such conflicts result in group break‐ups, and are therefore a fundamental driver of fusion–fission group dynamics. Yet, a paucity of empirical work hampers our understanding of how adaptive evolution has shaped plasticity in collective behaviours that promote and maintain fusion–fission dynamics. Using movement data from GPS‐collared bison, we found that individuals constantly associated with other animals possessing different spatial knowledge, and both personal and conspecific information influenced an individual's patch choice decisions. During conflict situations, bison used group familiarity coupled with their knowledge of local foraging options and recently sampled resource quality when deciding to follow or leave a group – a tactic that led to energy‐rewarding movements. Natural selection has shaped collective behaviours for coping with social conflicts and resource heterogeneity, which maintain fusion–fission dynamics and play an essential role in animal distribution.     

Mother–offspring distances reflect sex differences in fine‐scale genetic structure of eastern grey kangaroos

Natal dispersal affects life history and population biology and causes gene flow. In mammals, dispersal is usually male‐biased so that females tend to be philopatric and surrounded by matrilineal kin, which may lead to preferential associations among female kin. Here we combine genetic analyses and behavioral observations to investigate spatial genetic structure and sex‐biased dispersal patterns in a high‐density population of mammals showing fission–fusion group dynamics. We studied eastern grey kangaroos (Macropus giganteus) over 2 years at Wilsons Promontory National Park, Australia, and found weak fine‐scale genetic structure among adult females in both years but no structure among adult males. Immature male kangaroos moved away from their mothers at 18–25 months of age, while immature females remained near their mothers until older. A higher proportion of male (34%) than female (6%) subadults and young adults were observed to disperse, although median distances of detected dispersals were similar for both sexes. Adult females had overlapping ranges that were far wider than the maximum extent of spatial genetic structure found. Female kangaroos, although weakly philopatric, mostly encounter nonrelatives in fission–fusion groups at high density, and therefore kinship is unlikely to strongly affect sociality.     

The importance of fission–fusion social group dynamics in birds

Almost all animal social groups show some form of fission–fusion dynamics, whereby group membership is not spatio‐temporally stable. These dynamics have major implications at both population and individual levels, exerting an important influence on patterns of social behaviour, information transfer and epidemiology. However, fission–fusion dynamics in birds have received relatively little attention. We review the existing evidence for fission–fusion sociality in birds alongside a more general explanation of the social and ecological processes that may drive fission–fusion dynamics. Through a combination of recent methodological developments and novel technologies with well‐established areas of ornithological research, avian systems offer great potential to further our understanding of fission–fusion social systems and the consequences they have at an individual and population level. In particular, investigating the interaction between social structure and environmental covariates can promote a deeper understanding of the evolution of social behaviour and the adaptive value of group living, as well as having important consequences for applied research.     

Relationship quality affects fission decisions in wild spider monkeys (Ateles geoffroyi)

Fission–fusion dynamics are thought to be mainly a response to differential availability of food resources. However, social factors may also play a role. Here, we examined whether the quality of social relationships between group members affects fission decisions. During 21 months, we collected data on social interactions and fission events of 22 spider monkeys (Ateles geoffroyi) living in a community in the protected area of Otoch Ma'ax Yetel Kooh, Yucatan, Mexico. By entering seven indexes of social interactions into a principal component analysis, we obtained three components of relationship quality, which we labelled “compatibility,” “value” and “insecurity” given the relative loadings of the indexes. Our results showed that individuals were more likely to fission into the same subgroup with community members with whom they shared higher levels of compatibility and value and lower levels of insecurity. In addition, individuals preferred to fission into the same subgroup with same‐sex group members, as expected based on what is known for the species. Our findings highlight the role of social factors in fission decisions. Adjustments in subgroup size are based on multifaceted social preferences, incorporating previously unexamined aspects of relationship quality, which are independent from overall levels of affiliative interactions.     

Niche overlap and diet composition of three sympatric coastal dolphin species in the southwest Atlantic Ocean

Sympatric species are expected to differ in ecological requirements to minimize niche overlap and avoid competition. Here we assess the trophic interactions among three coexisting dolphin species from southern Brazil: the franciscana dolphin (Pontoporia blainvillei), the Guiana dolphin (Sotalia guianensis), and the Lahille's bottlenose dolphin (Tursiops truncatus gephyreus). We evaluated temporal variation in carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope values of bone collagen to examine potential dietary shifts resulting from increased fishing activity over the past three decades. We estimated the degree of niche overlap among these species and the contribution of potential prey sources to their diet. δ¹⁵N values were consistent among species and across years, while δ¹³C values increased for Guiana dolphins and decreased for bottlenose dolphins, suggesting changes in diet and/or foraging habitats through time. The similar δ¹³C and δ¹⁵N values and the high niche overlap between Guiana and bottlenose dolphins indicate that these species are primarily feeding on demersal prey. The franciscana diet is primarily composed of pelagic prey, resulting in a lower niche overlap in comparison with the other dolphin species. Our study provides further information about the foraging ecology of this unique dolphin community in southern Brazil with implications for its management and conservation.     

Collective decision‐making and fission–fusion dynamics: a conceptual framework

Sociality exists in an extraordinary range of ecological settings. For individuals to accrue the benefits associated with social interactions, they are required to maintain a degree of spatial and temporal coordination in their activities, and make collective decisions. Such coordination and decision‐making has been the focus of much recent research. However, efforts largely have been directed toward understanding patterns of collective behaviour in relatively stable and cohesive groups. Less well understood is how fission–fusion dynamics mediate the process and outcome of collective decisions making. Here, we aim to apply established concepts and knowledge to highlight the implications of fission–fusion dynamics for collective decisions, presenting a conceptual framework based on the outcome of a small‐group discussion INCORE meeting (funded by the European Community's Sixth Framework Programme). First, we discuss how the degree of uncertainty in the environment shapes social flexibility and therefore the types of decisions individuals make in different social settings. Second, we propose that the quality of social relationships and the energetic needs of each individual influence fission decisions. Third, we explore how these factors affect the probability of individuals to fuse. Fourth, we discuss how group size and fission–fusion dynamics may affect communication processes between individuals at a local or global scale to reach a consensus or to fission. Finally, we offer a number of suggestions for future research, capturing emerging ideas and concepts on the interaction between collective decisions and fission–fusion dynamics.     

Fine‐scale habitat use in Indo‐Pacific humpback dolphins,Sousa chinensis,may be more influenced by fish rather than vessels in the Pearl River Estuary,China

Shifts in habitat use and distribution patterns in dolphins are often concerns that can result from habitat degradation. We investigated how potential changes to a habitat from human activity may alter dolphin distributions within Lingding Bay in the Pearl River Estuary, China, by studying the relationship between fish choruses, vessel presence and Indo‐Pacific humpback dolphin (Sousa chinensis) detection rates. Analyses revealed temporal and spatial variation within fish choruses, vessel presence and dolphin detection rates. After accounting for any temporal autocorrelation, correlations between fish choruses and dolphin detection rates were also found; however, no relationship between fish choruses and vessel presence or dolphin detection rates and vessel presence were observed. Furthermore, fewer dolphins were detected at sites where fish activity was less intense. Thus fish activity, rather than vessels, may be a key factor influencing the distribution of the dolphins within the estuary. These findings emphasize the risk of potential shifts in habitat use for Indo‐Pacific humpback dolphins due to detrimental changes to prey availability and dolphin feeding grounds from human activity, such as overfishing and coastal developments, within the estuary. This is a critical conservation issue for this dolphin population that is facing intense anthropogenic pressure.     

Herd composition,kinship and fission–fusion social dynamics among wild giraffe

A variety of social systems have evolved as a consequence of competition and cooperation among individuals. Giraffe (Giraffa camelopardalis sp.) societies are an anomaly because the dearth of long‐term data has produced two polar perspectives: a loose amalgamation of non‐bonded individuals that sometimes coalesce into a herd and a structured social system with a fission–fusion process modifying herd composition within a community. We analysed 34 years of data collected from a population of Thornicroft's giraffe (G. c. thornicrofti, Lydekker 1911) residing in South Luangwa, Zambia, to establish the nature of giraffe society. Our sample consisted of 52 individually recognized animals. We found that giraffe herd composition is based upon long‐term social associations that often reflect kinship, with close relatives significantly more likely than non‐relatives to establish herds. Mother/offspring dyads had the strongest associations, which persisted for years. Giraffe live in a complex society characterized by marked flexibility in herd size, with about 25% of the variance in herd composition owing to kinship and sex. We suggest that giraffe herds share many characteristics of fission–fusion social systems and propose that sophisticated communication systems are a crucial component regulating subgroup dynamics.     

Breathing synchrony in franciscana (Pontoporia blainvillei) and Guiana dolphins (Sotalia guianensis) in Southern Brazil

Synchronous breathing may be a useful proxy for studying other, and perhaps more complex, aspects of cetacean behavior. Here we describe a study conducted in Babitonga Bay, southern Brazil, where we investigated the synchrony of breathing in two small populations of franciscana (Pontoporia blainvillei) and Guiana dolphins (Sotalia guianensis). The bay is affected by different sources of anthropogenic disturbances, such as boat activity and point‐source pollution. We assumed breathing synchrony to be the inverse of the time between breathing surfacing displays of dolphins within a swimming pair, which we refer to as lag. The relationship between lag and anthropogenic and animal‐related variables was evaluated with generalized additive models. For franciscana dolphins, breathing synchrony was only positively related to the proximity between animals. Breathing synchrony in Guiana dolphins was positively related to both the proximity between animals and to group size, and varied significantly depending on the research boat used. Proximal characteristics (i.e., of individuals or of the group) of these dolphin species seem to be more related to the synchronization in breathing than are the environmental variables assessed here. Results presented expand the current knowledge of these two dolphin species and provide general insights into the breathing synchrony for cetaceans.     

Long‐term social structure of a resident community of Atlantic spotted dolphins,Stenella Frontalis,in the Bahamas 1991–2002

Long‐term social structure data on small delphinids is lacking for most species except the bottlenose dolphin. This study describes the long‐term social structure of one community of Atlantic spotted dolphins, Stenella frontalis, divided into three social clusters. Data from 12 yr were analyzed using SOCPROG 2.3. Coefficients of association (CoA) were calculated using the half‐weight index. The overall mean community CoA ranged from 0.09 to 0.12. Temporal analyses and mantel tests revealed significant differences between sex class associations due to high male‐male CoA (0.12–0.23) compared to female‐female and mixed sex CoA (0.08–0.10). Female associations were strongly influenced by reproductive status, calf care, and social familiarity, but not by age class. Male associations were strongly influenced by age, access to females, and alliance formation. Males showed two levels of alliance formation, long‐term first order pairs/trios (CoA 0.70–1.00) and shorter‐term second order alliances between two or more first order alliances (CoA 0.45–0.69), and a possible third level during interspecies interactions. Mating strategies, sex, and cluster formation shaped the social structure in this spotted dolphin community. Similar to many bottlenose dolphin studies, long‐term affiliations for spotted dolphins were correlated with age, sex, and reproductive status.