Affect Group Size Research Articles

Context-dependent group size: effects of population density, habitat, and season

AbstractGroup size can vary in relation to population density, habitat, and season. Habitat and season may also interact with population density and affect group size through varying foraging benefits of social aggregation in different ecological contexts. We tested the hypothesis that group size varies across ecological contexts, including population density, habitat type, and season, for woodland caribou (Rangifer tarandus) in ten herds over 25 years in Newfoundland, Canada. We predicted that group size would increase as a function of population density. Based on the foraging benefits of social aggregation, we predicted larger groups as habitat openness increased because open areas tend to have higher quality foraging resources. We predicted larger groups during winter when foraging resources are covered in snow because caribou and other social animals exploit social information about the location of foraging resources. In contrast to our prediction, group size decreased as a function of population density. In support of our prediction, group size was larger in winter than calving and summer, and we found that group size increased with habitat openness in some, but not all, cases. Patterns of animal grouping are context-dependent and the additive effect of different ecological contexts on variation in group size informs our understanding of the implicit trade-offs between competition, predation risk, and profitability of forage.

Dynamic visual noise promotes social attraction, but does not affect group size preference, in a shoaling fish

Gathering information from the environment allows animals to make informed behavioural decisions, with individuals sampling information either privately or via social cues from group members. Environmental noise, however, may disrupt the ability of animals to gather information in their environment. Therefore, social interactions could be disrupted by environmental noise, or adapted to mitigate the costs associated with compromised perception in noisy environments. Here we tested whether the presence of water caustics, a natural form of visual noise prevalent within shallow aquatic habitats, affects the social decisions of shoaling fish. In a classic experimental paradigm, individual three-spined sticklebacks, Gasterosteus aculeatus, were given a choice between two conspecific shoals differing in size while immersed in different levels of visual noise. Fish showed greater social attraction towards conspecifics as noise levels increased, measured by a reduction in the distance they maintained from others. While sticklebacks only showed clear preferences for groups that had at least four more members than another group, there was no evidence that visual noise level influenced fish's preference to associate with these larger shoals, nor did it change the preferences for other shoal sizes. Our findings suggest that, at the group sizes tested, visual noise increases individual stickleback's social attraction towards conspecifics, but does not affect their decision to associate with the larger of two groups. Such changes to social behaviour may reflect the consequences of having a reduced ability to sample private, but not social, information in noisy environments.

Machine Learning Data Imputation and Prediction of Foraging Group Size in a Kleptoparasitic Spider

Cost–benefit analysis is widely used to elucidate the association between foraging group size and resource size. Despite advances in the development of theoretical frameworks, however, the empirical systems used for testing are hindered by the vagaries of field surveys and incomplete data. This study developed the three approaches to data imputation based on machine learning (ML) algorithms with the aim of rescuing valuable field data. Using 163 host spider webs (132 complete data and 31 incomplete data), our results indicated that the data imputation based on random forest algorithm outperformed classification and regression trees, the k-nearest neighbor, and other conventional approaches (Wilcoxon signed-rank test and correlation difference have p-value from < 0.001–0.030). We then used rescued data based on a natural system involving kleptoparasitic spiders from Taiwan and Vietnam (Argyrodes miniaceus, Theridiidae) to test the occurrence and group size of kleptoparasites in natural populations. Our partial least-squares path modelling (PLS-PM) results demonstrated that the size of the host web (T = 6.890, p = 0.000) is a significant feature affecting group size. The resource size (T = 2.590, p = 0.010) and the microclimate (T = 3.230, p = 0.001) are significant features affecting the presence of kleptoparasites. The test of conformation of group size distribution to the ideal free distribution (IFD) model revealed that predictions pertaining to per-capita resource size were underestimated (bootstrap resampling mean slopes <IFD predicted slopes, p < 0.001). These findings highlight the importance of applying appropriate ML methods to the handling of missing field data.

The components of directional and disruptive selection in heterogeneous group-structured populations

We derive how directional and disruptive selection operate on scalar traits in a heterogeneous group-structured population for a general class of models. In particular, we assume that each group in the population can be in one of a finite number of states, where states can affect group size and/or other environmental variables, at a given time. Using up to second-order perturbation expansions of the invasion fitness of a mutant allele, we derive expressions for the directional and disruptive selection coefficients, which are sufficient to classify the singular strategies of adaptive dynamics. These expressions include first- and second-order perturbations of individual fitness (expected number of settled offspring produced by an individual, possibly including self through survival); the first-order perturbation of the stationary distribution of mutants (derived here explicitly for the first time); the first-order perturbation of pairwise relatedness; and reproductive values, pairwise and three-way relatedness, and stationary distribution of mutants, each evaluated under neutrality. We introduce the concept of individual k-fitness (defined as the expected number of settled offspring of an individual for which k-1 randomly chosen neighbors are lineage members) and show its usefulness for calculating relatedness and its perturbation. We then demonstrate that the directional and disruptive selection coefficients can be expressed in terms individual k-fitnesses with k=1,2,3 only. This representation has two important benefits. First, it allows for a significant reduction in the dimensions of the system of equations describing the mutant dynamics that needs to be solved to evaluate explicitly the two selection coefficients. Second, it leads to a biologically meaningful interpretation of their components. As an application of our methodology, we analyze directional and disruptive selection in a lottery model with either hard or soft selection and show that many previous results about selection in group-structured populations can be reproduced as special cases of our model.

Truncated power-law distribution of group sizes in antelope

Abstract Quantifying and understanding group size distributions can be useful for understanding group behaviour in animal populations. We analysed group size data of the blackbuck, Antilope cervicapra, from six different field sites to estimate the group size distribution of this antelope. We used likelihood based methods (AICs and likelihood ratios) to show that an exponentially truncated power law is the distribution that best describes blackbuck group data, outperforming a simple power-law, an exponential distribution, and a lognormal distribution. Our results show that distribution parameters can be used to draw novel insights regarding group dynamics, and we demonstrate this by investigating how habitat openness affects group size distributions.

Seasonal changes and the influence of tree species and ambient temperature on the fission-fusion dynamics of tree-roosting bats

Many species of bats live in dynamic associations in which individuals aggregate or split into smaller or larger groups, moving between roost sites, but factors influencing group size and roost switching decisions in bats living in fission-fusion societies are poorly understood. To test the extent to which fission-fusion dynamics are species-specific and shaped by environmental factors, we radio-tracked two closely related species: noctule bats Nyctalus noctula and Leisler’s bats N. leisleri in well-preserved, old-growth stands of the Białowieża Forest, where roost availability and diversity is high and bats can potentially freely exhibit fission-fusion behaviour. We estimated the number of bats per roost and noted the tree species selected as roost sites. We evaluated the influence of ambient temperature and roost tree species on bat group size and the process of roost switching, and their seasonal component. Our results revealed that bat group size showed species-specific differences in seasonal variation, relationship to ambient temperature and tree species. The number of noctule bats emerging from a roost was relatively stable with a weak tendency to decrease during the breeding season while the number of Leisler’s bats emerging from a roost increased when the juveniles started to be volant. Group size of noctule bats increased with ambient temperature. Leisler’s bats formed smaller colonies in oaks and ashes than in other tree species. Roost switching rate was associated with ambient temperature and tree species selected by the bats. The bats changed roosts more frequently when the minimum ambient temperature increased, and less often in ashes than in oaks and other tree species. Our results show that bat group size and roost switching behaviour are shaped by both environmental factors and the stage of reproduction.Significance statementEnvironmental factors may affect group size and roost switching behaviour of bats forming fission-fusion associations, but this is expected to vary depending on body size, environmental conditions, as well as seasonally. Few studies have explored this behaviour in tree-dwelling bats and examined how it differs between species. Here, we studied the group size dynamics and the process of roost switching in closely related bat species living in well-preserved, European old-growth forests, for which the knowledge of these patterns is particularly poor. We demonstrate for the first time that bat group size and roost switching rate relates to roost tree species selected by bats, the ambient temperature and varies seasonally, and that these effects are species-specific. The results point out the importance of the availability of suitable roost trees providing a range of microclimatic conditions for maintaining efficient fission-fusion dynamics throughout the reproductive period.

Use of social thermoregulation fluctuates with mast seeding and reproduction in a pulsed resource consumer

Edible dormice (Glis glis) can remain entirely solitary but frequently share sleeping sites with conspecifics in groups of up to 16 adults and yearlings. Here, we analysed grouping behaviour of 4564 marked individuals, captured in a 13-year study in nest boxes in a deciduous forest. We aimed to clarify (i) whether social thermoregulation is the primary cause for group formation and (ii) which factors affect group size and composition. Dormice temporarily formed both mixed and single-sex groups in response to acute cold ambient temperatures, especially those individuals with small body mass. Thus, thermoregulatory huddling appears to be the driving force for group formation in this species. Huddling was avoided—except for conditions of severe cold load—in years of full mast seeding, which is associated with reproduction and high foraging activity. Almost all females remained solitary during reproduction and lactation. Hence, entire populations of dormice switched between predominantly solitary lives in reproductive years to social behaviour in non-reproductive years. Non-social behaviour pointed to costs of huddling in terms of competition for local food resources even when food is generally abundant. The impact of competition was mitigated by a sex ratio that was biased towards males, which avoids sharing of food resources with related females that have extremely high energy demands during lactation. Importantly, dormice preferentially huddled in male-biased groups with litter mates from previous years. The fraction of related individuals increased with group size. Hence, group composition partly offsets the costs of shared food resources via indirect fitness benefits.

Allee effects drive the coevolution of cooperation and group size in high reproductive skew groups

Abstract The evolution of cooperation between conspecifics is a fundamental evolutionary puzzle, with much work focusing on the evolution of cooperative breeding. Surprisingly, although we expect cooperation to affect the population structures in which individuals interact, most studies fail to allow cooperation and population structure to coevolve. Here, we build two models containing group-level Allee effects (positive density dependence at low group sizes) to study the coevolution of cooperation and group size. Group-level Allee effects, although common in cooperatively breeding species, remain understudied for their evolutionary implications. We find that a trait that affects group size can cause increased cooperation to be favored evolutionarily even in a group with complete reproductive skew. In particular, we find a single evolutionarily stable attractor in our model corresponding to moderate helpfulness and group size. In general, our results demonstrate that, even in groups with complete reproductive skew, Allee effects can be important for the evolution of cooperation and that the evolution of cooperation may be closely linked to the evolution of group size. Further, our model matches empirical data in African wild dogs (Lycaon pictus), suggesting that it may have an application in understanding social evolution in this endangered species.

Fission-fusion dynamics of a megaherbivore are driven by ecological, anthropogenic, temporal, and social factors.

Fission-fusion dynamics hypothetically enable animals to exploit dispersed and ephemeral food resources while minimizing predation risk. Disentangling factors affecting group size and composition of fission-fusion species facilitates their management and conservation. We used a 6-year data set of 2888 group formations of Masai giraffes in Tanzania to investigate determinants of social group size and structure. We tested whether ecological (lion density, vegetation structure, and prevalence of primary forage plants), anthropogenic (proximity to human settlements), temporal (rainy or dry season), and social (local giraffe density, adult sex ratio, and proportion of calves) factors explained variation in group size and sex- and age-class composition. Food availability rather than predation risk mediated grouping dynamics of adult giraffes, while predation risk was the most important factor influencing congregations with calves. Smallest group sizes occurred during the food-limiting dry season. Where predation risk was greatest, groups with calves were in bushlands more than in open grasslands, but the groups were smaller in size, suggesting mothers adopted a strategy of hiding calves rather than a predator-detection-and-dilution strategy. Groups with calves also were farther from towns but closer to traditional human compounds (bomas). This may be due to lower predator densities, and thus reduced calf predation risk, near bomas but higher human disturbance near towns. Sex- and age-based differences in habitat use reflected nursing mothers' need for high-quality forage while also protecting their young from predation. Our results have implications for conservation and management of giraffes and other large-bodied, herd-forming ungulates in heterogeneous environments subject to anthropogenic threats.

Climatic variation across a latitudinal gradient affect phenology and group size, but not social complexity in small carpenter bees

Greater social complexity at lower latitudes has been observed in a variety of arthropods from termites to spiders. Social behavior in the small carpenter bees, Ceratina, has been shown to vary widely both between species and across geographic range. Our goal was to determine how social plasticity of three populations of Ceratina species, C. calcarata and C. strenua, vary across a latitudinal gradient. The longer rearing season in the south did not result in two separate brood rearing periods, but instead increased brood production of a single brood with a higher female sex bias. The social structure of nests remained stable across both species’ ranges: mothers exhibit prolonged parental care and worker dwarf eldest daughters occur among populations and species. This is the first report of worker daughters in C. strenua. The ubiquity of worker daughter production in eastern North American Ceratina suggests that factors outside of climate underlie the early division of labor between the reproductive mother and worker dwarf eldest daughter.

Group sizes of giraffes in Kenya: the influence of habitat, predation and the age and sex of individuals

AbstractGiraffe group sizes appear to vary in response to localized ecological and environmental factors, but there has been little investigation of how social factors or predation risk affect group size in giraffes. We studied two adjacent, enclosed populations of Rothschild's giraffes in Kenya, and used 591 records of groups to determine the relative influence of a series of variables on group size. One population was free from any risk of predation, while the other area contained a high density of lions. Mean group size was smaller in the population with lions, but a series of Generalized Linear Mixed Models accounting for habitat and age/sex class of individuals showed that the presence of high numbers of juveniles in the area free from lions artificially inflated group sizes. Removing juveniles from the analysis showed that contrary to the existing creche hypothesis, adult females were found in smaller groups when they had calves. We found no evidence that predation risk influenced grouping behaviour. Rather, recruitment and habitat type had a stronger influence on group sizes, but the results were complex and varied between different age and sex classes of individual. We conclude that predation is not an important driver of giraffe grouping, and that further research is necessary to understand the complex behaviour and ecology of this prominent yet understudied species.

Individual consistency and context dependence in group-size preference of Eurasian perch

Many fish spend a large part of their life in groups. The size of the group influences potential costs and benefits of group living, and depending on context a fish may prefer different group sizes. Group-size preference may also depend on personality, with social individuals expected to prefer larger groups than asocial fish. This study investigates context-dependent group size preference in two populations of a highly social fish, young of the year Eurasian perch. The perch were given a choice between a group of two and a group of eight conspecifics under three different situations: the small group was feeding, the small group had access to shelter, and a control treatment with no extra stimuli. In general, the perch associated more with the large group, but significantly less so during the food treatment. Perceived access to shelter did not affect group size preference compared to the control treatment. Consistent individual differences in social attraction were found within each context, but not among all contexts. Also, an individual’s sociability did not correlate with its degree of boldness, indicating a lack of a behavioural syndrome between the two personality traits in the studied populations. The results highlight the importance of considering environmental context when studying social behaviour in obligate social fish, and show the complexity of the concept of sociability as a personality trait by demonstrating context dependence in individual consistency in social behaviour.

Responses of two sympatric species of peccaries (Tayassu pecari and Pecari tajacu) to hunting in Calakmul, Mexico

The white-lipped peccary ( Tayassu pecari) and the collared peccary ( Pecari tajacu) are social ungulates of great ecological and economic importance. Both species represent important sources of animal protein in the diet of indigenous and rural communities in Neotropical forests, which has contributed to reducing their populations throughout their geographic distribution. It is still not well understood how the social ecology of peccaries is affected by hunting. This study analyzed the relationship between hunting and group size, group composition, daily activity patterns, and the relative abundance of the two peccary species. Camera-traps and direct observation were used from February 2014 through February 2015 in two sites differing in their degree of protection and hunting pressure: the first was Calakmul Biosphere Reserve (a site with no hunting) and the second was the community of Nuevo Becal (a hunting site). Our analyses suggest that hunting negatively affects group size, structure, composition and the relative abundance of both peccary species. We propose that management and conservation plans, particularly for key resources such as water, and monitoring and hunting controls in the habitats of both peccary species, might ensure their survival in the Calakmul region.

What Ecological and Anthropogenic Factors Affect Group Size in White‐lipped Peccaries (Tayassu pecari)?

AbstractGroup living among ungulates has evolved mainly in species living in open habitats, such as grasslands and savannas, whereas in the forest, few ungulate species form groups and these tend to be small. Therefore, the white‐lipped peccary (Tayassu pecari), a Neotropical ungulate listed as Vulnerable by the IUCN, represents an almost unique social occurrence as it lives in large and cohesive groups, yet it inhabits dense tropical forests. Large variations in group sizes have been observed throughout the species range, with reports of herds with less than 10 to around 300 individuals. We examined factors that might cause variation in group size in white‐lipped peccary, including ecological and anthropogenic variables. We conducted an extensive literature review and used our original data to compile information on white‐lipped peccary's group size across its range. We built models to quantitate generalizations for group sizes distinguishing data from areas with high human influence, and areas that have not been significantly disturbed by humans for at least the last 20 years. We found that white‐lipped peccary's group size for all sites was most strongly predicted by a combination of the distances to the nearest human settlement and rainfall and its seasonality. Results from the undisturbed sites indicated that group size is positively influenced by rainfall. Our results contribute to understand why group size varies in different environments that are subjected to different ecological and human conditions. Information on these relationships is a key to advance our understanding of the socio‐ecological strategies of animal species living in groups.

Genotype and spatial structure shape pathogen dispersal and disease dynamics at small spatial scales.

Many devastating pathogens are passively dispersed, and their epidemics are characterized by variation that is typically attributed to environmental factors. Here, by combining laboratory inoculations with wind tunnel and field trials using the wind-dispersed pathogen Podosphaera plantaginis, we demonstrated striking genetic variation affecting the unexplored microscale (< 2 m) of epidemics. Recipient and source host genotypes, as well as pathogen strain, explained a large fraction of variation in the three key dispersal phases: departure, movement, and settlement. Moreover, we found genotypic variation affecting group size of the pathogen dispersal unit, ultimately resulting in increased disease development on hosts close to the infection source. Together, our results show that genotypic variation may generate considerable variation in the rate of disease spread through space and time with disease hotspots emerging around initial foci. Furthermore, the extent of genetic variation affecting the entire dispersal process confirms that these traits may be targeted by selection.

Density-Dependent Effects on Group Size Are Sex-Specific in a Gregarious Ungulate

Density dependence can have marked effects on social behaviors such as group size. We tested whether changes in population density of a large herbivore (elk, Cervus canadensis) affected sex-specific group size and whether the response was density- or frequency-dependent. We quantified the probability and strength of changes in group sizes and dispersion as population density changed for each sex. We used group size data from a population of elk in Manitoba, Canada, that was experimentally reduced from 1.20 to 0.67 elk/km2 between 2002 and 2009. Our results indicated that functional responses of group size to population density are sex-specific. Females showed a positive density-dependent response in group size at population densities ≥0.70 elk/km2 and we found evidence for a minimum group size at population density ≤0.70 elk/km2. Changes in male group size were also density-dependent; however, the strength of the relationship was lower than for females. Density dependence in male group size was predominantly a result of fusion of solitary males into larger groups, rather than fusion among existing groups. Our study revealed that density affects group size of a large herbivore differently between males and females, which has important implications for the benefits e.g., alleviating predation risk, and costs of social behaviors e.g., competition for resources and mates, and intra-specific pathogen transmission.

Fission–Fusion Behavior in Yunnan Snub-Nosed Monkeys (Rhinopithecus bieti) in Yunnan, China

Several species of mammals live in relatively large social groups that temporarily fission or subgroup in response to changes in food availability, predation risk, foraging strategies, and mating competition. Although the dynamics of subgrouping are not well understood, evidence of facultative fission–fusion behavior in species that generally exhibit a highly cohesive group structure may help to clarify the set of ecological and social factors that constrain or affect group size in primates. We here examined patterns of subgrouping in Yunnan snub-nosed monkeys (Rhinopithecus bieti) inhabiting the Baimaxueshan National Nature Reserve in Yunnan, China. Rhinopithecus bieti lives in a large multilevel society consisting of 6–41 one-male, multifemale units (OMUs) and one all-male unit (AMU). Over a 5-yr period from 2003 through 2008, we tracked a group of >450 Rhinopithecus bieti. We recorded the location in the group’s home range where fission and fusion events occurred, the size and number of subgroups, days spent in subgroups, and food availability, and monitored predator sightings. The results indicate that the focal group underwent12 group fission events, all of which occurred between mid-June and July, and all in the same area of the group’s range. During these fissioning events, the AMU also showed subgrouping behavior. In all cases, the presence of bamboo shoots, an important seasonal component of the snub-nosed monkey diet, appeared to trigger subgrouping behavior. The subgroups reunited in other parts of their range after a period of 2–11 d. We found no evidence that the presence of predators had a direct affect on subgrouping behavior in Rhinopithecus bieti, as proposed in previous studies.

Boundary nature induces greater group size and group density in habitat edges: an agent‐based model revealed

AbstractApart from the amount and distribution of food resources and predation pressure, we sought to clarify habitat spatial features that affect group size and group density. To simulate animal groups interacting with one another, we made an agent‐based model in a computational space. Assuming no predation pressure, we change the resource conditions between the habitat (comprising ‘interior’ and ‘edges’) and the ‘exterior’. The results are as follows: when the group density is high throughout the habitat, large groups tend to frequent the edges (i.e., the boundaries between the interior and exterior) even if food resources are scarce in the exterior. Additionally, when the group density is high and the group size variation is small, both group size and group density increase at the edges. These findings are discussed with reference to primates, particularly the Japanese macaque (Macaca fuscata) as an example of group‐living animal.

It takes grouping and cooperation to get sociality

Cooperation and grouping are regularly studied as separate traits. The evolution of sociality however requires both that individuals get together in groups and that they cooperate within them. Because the level of cooperation can influence selection for group size, and vice versa, it is worth studying how these traits coevolve. Using a generally applicable two-trait optimization approach, we provide analytical solutions for three specific models. These solutions describe how cooperative associations of non-relatives evolve, and predict how large and how cooperative they will be. The analytical solutions help understand how changes in parameter values, such as the group carrying capacity and the costs of cooperation, affect group size and the level of cooperation in equilibrium. Although the analytical model makes a few simplifying assumptions—populations are assumed to be monomorphic for grouping as well as for cooperative tendencies, and group size is assumed to be deterministic—simulations show that its predictions are matched quite closely by results for settings where these assumptions do not hold.

CnrN regulates Dictyostelium group size using a counting factor-independent mechanism

One of the simplest examples of a complex behavior is the aggregation of solitary Dictyostelium discoideum amoebae to form a 20,000-cell fruiting body. A field of starving amoebae first breaks up into territories. In each territory, the cells form a spider-like pattern of streams of cells. As part of a negative feedback loop, counting factor (CF), a secreted protein complex whose concentration increases with the size of the stream, prevents over-sized fruiting bodies from being formed by increasing cell motility and decreasing cell-cell adhesion, which causes the breakup of excessively large streams. Cells lacking the phosphatase CnrN (cnrN- cells) form small aggregation territories and few streams.1 In this report, we present computer simulations that suggest that in the absence of stream formation, CF should be unable to affect group size. As predicted, cnrN- group size is insensitive to the addition or depletion of CF. Together, the data indicate that CnrN regulates group size by regulating both the break-up of a field of cells into aggregation territories and stream formation during development, and that CnrN-mediated and CF-mediated group size regulation use different mechanisms.