Group size versus territory size in group-living badgers: a large-sample field test of the Resource Dispersion Hypothesis

Badgers ( Meles meles ) have been the focus for the development of a pervasive model of social grouping behaviour, relevant to a number of carnivore species and other taxonomic groups – the Resource Dispersion Hypothesis (RDH). The RDH hypothesises that the dispersion and richness of resources in the environment provide a passive mechanism for the formation of groups, even without any direct benefits of group living. However, few studies have tested the RDH in the field. The principal prediction is that, as opposed to enlargement of territory sizes to accommodate more members, territory size (TS) is independent of group size (GS). Instead, TS is determined by the spatial dispersion of resources, while GS is independently determined by the richness of those resources. However, these predictions provide only weak correlative tests, especially in non-experimental field studies. The first predicts an absence of correlation and is therefore prone to Type II error, especially given the small sample sizes and errors in estimating TS and GS of mammals in the field. We tested for independence of territory size and group size in all years with available data since the beginning of the long-term badger study in Wytham Woods in 1974. We used two methods of TS estimation, a sequential Bonferroni technique to adjust for multiple inference tests, a combined analysis and an analysis with pooled data. This prediction of the RDH could not be rejected on the basis of any of these analyses. Given this evidence that other processes are independently determining group size and territory size, further predictions of the RDH will be worth investigating in considerable detail.     

Group size correlates with territory size in European badgers: implications for the resource dispersion hypothesis?

The resource dispersion hypothesis (RDH) predicts that resource heterogeneity can act as a passive cause of group‐living in social carnivores and potentially many other species. One central prediction of the RDH is that territory size and group size are not related, as they are determined by resource dispersion and quality, respectively. In this study we investigated the relationship between territory size, group size and group composition in the European badger, a non‐cooperative social mustelid whose behavioural ecology was central to the development of the RDH. Using data from a long‐term study in the UK, we found that territory size and group size were positively related, contradicting a core prediction of the RDH. Furthermore, territory size was more strongly correlated with the number of adult males in the group than to the number of females or total group size. This result suggests that male badgers may have a more important role in territoriality and receive greater benefits from territory enlargement. These findings are consistent with the predictions of the anti‐kleptogamy hypothesis, and suggest that badger territorial and social behaviour is not purely driven by resource dispersion, but may also be associated with breeding behaviour, as in other mustelids.     

Influence of Fruit Availability on the Fission–Fusion Dynamics of Spider Monkeys (<Emphasis Type="Italic">Ateles geoffroyi</Emphasis>)

Socioecological theory proposes that the flexibility in grouping patterns afforded by fission–fusion dynamics allows animals to cope with spatiotemporal variability in food abundance. We investigate the influence of fruit tree abundance and foraging environment heterogeneity on fission–fusion dynamics in a group of spider monkeys (Ateles geoffroyi) in the Yucatan peninsula, Mexico. We collected 1300 h of behavioral data and 23 samples of biweekly ecological data from August 2009 to July 2010. We measured fission–fusion dynamics through the temporal variation in the size and composition of subgroups, the spatial dispersion within and between subgroups, and the frequency of fissions and fusions. We measured habitat-wide food abundance of preferred species, including two that differ greatly in their relative abundance: Brosimum alicastrum (a hyperabundant resource) and Ficus spp. (a not so abundant resource but often represented by large trees). We evaluated the foraging environment heterogeneity through the variance in the number of trees with fruit between species. Our results show that, although habitat-wide food abundance is important, the availability of key resources strongly influences the spider monkeys’ fission–fusion dynamics. When there was a high abundance of fruit of Brosimum, subgroups tended to be more stable, smaller, and mixed sex, and their members remained close. In contrast, when Brosimum trees with fruit were scarce, females often formed large, more fluid and dispersed subgroups. Foraging environment heterogeneity had a positive effect on within-subgroup spatial dispersion and rates of fission and fusion. The complex relationships we have uncovered suggest that the flexibility afforded by fission–fusion dynamics is an adaptation to highly variable foraging environments.     

Home range,activity and sociality of a top predator,the dingo: a test of the Resource Dispersion Hypothesis

The idea that groups of individuals may develop around resource patches led to the formulation of the Resource Dispersion Hypothesis (RDH). We tested the predictions of the RDH, within a quasi‐experimental framework, using Australia’s largest terrestrial predator, the dingo Canis lupus dingo. Average dingo group sizes were higher in areas with abundant focal food sources around two mine sites compared with those in more distant areas. This supports the notion that resource richness favours larger group size, consistent with the RDH. Irrespective of season or sex, average home range estimates and daily activity for dingoes around the mine sites were significantly less than for dingoes that lived well away. Assuming that a territory is the defended part of the home range and that territory size is correlated with home range size, consistent with the RDH, the spatial dispersion of food patches therefore determined territory size for dingoes in our study. However, although sample size was small, some dingoes that accessed the supplementary food resource at the mines also spent a large proportion of their time away, suggesting a breakdown of territorial defence around the focal food resource. This, in combination with the large variation in home range size among dingoes that accessed the same supplementary food resource, limits the predictive capabilities of the RDH for this species. We hypothesize that constraints on exclusive home range occupancy will arise if a surfeit of food resources (in excess of requirements for homeostasis) is available in a small area, and that this will have further effects on access to mates and social structure. We present a conceptual model of facultative territorial defence where focal resources are available to demonstrate our findings.     

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.     

The sociality of solitary foragers: a model based on resource dispersion

Under versions of what may broadly be called the Resource Dispersion Hypothesis (RDH) several authors have concluded that territory size and group size are limited respectively, and independently, by the dispersion and richness of patches of food. This paper presents a model that shows how the frequency distribution of resources available per unit time within a territory may permit the formation of groups even in the absence of any functional advantage to any individual from the presence of another. In this model, animals (called primary occupants) occupy territories containing sufficient resources to meet or exceed their requirements for a critical proportion of feeding periods. The availability of these resources is described in terms of their mean richness and their heterogeneity, and plots of these parameters indicate the circumstances within which individuals may share the minimum territory with the primary occupants. The model shows how, under plausible conditions of resource dispersion, a territory that provides almost total food security for two occupants could also provide, at no cost to the original occupants, substantial food security for an additional group member, even if it never used the same food patches as the originals. It is not therefore, as is sometimes supposed, a necessary condition for the RDH that members of a group often forage simultaneously in the same patch. Thus the model describes ecological circumstances whereby groups could evolve amongst species whose members neither forage communally, nor even meet frequently.     

Are fission–fusion dynamics consistent among populations? A large‐scale study with Cape buffalo

Fission–fusion dynamics allow animals to manage costs and benefits of group living by adjusting group size. The degree of intraspecific variation in fission–fusion dynamics across the geographical range is poorly known. During 2008–2016, 38 adult female Cape buffalo were equipped with GPS collars in three populations located in different protected areas (Gonarezhou National Park and Hwange National Park, Zimbabwe; Kruger National Park, South Africa) to investigate the patterns and environmental drivers of fission–fusion dynamics among populations. We estimated home range overlap and fission and fusion events between Cape buffalo dyads. We investigated the temporal dynamics of both events at daily and seasonal scales and examined the influence of habitat and distance to water on event location. Fission–fusion dynamics were generally consistent across populations: Fission and fusion periods lasted on average between less than one day and three days. However, we found seasonal differences in the underlying patterns of fission and fusion, which point out the likely influence of resource availability and distribution in time on group dynamics: During the wet season, Cape buffalo split and associated more frequently and were in the same or in a different subgroup for shorter periods. Cape buffalo subgroups were more likely to merge than to split in open areas located near water, but overall vegetation and distance to water were very poor predictors of where fission and fusion events occurred. This study is one of the first to quantify fission–fusion dynamics in a single species across several populations with a common methodology, thus robustly questioning the behavioral flexibility of fission–fusion dynamics among environments.     

Does the resource dispersion hypothesis explain group living?

The resource dispersion hypothesis (RDH) asserts that, if resources are heterogeneous in space or time, group living might be less costly than was previously thought, regardless of whether individuals gain direct benefits from group membership. The RDH was first proposed more than 20 years ago and has since accumulated considerable support. However, it is sometimes discredited because a priori tests of specific predictions are few, relevant variables have proved difficult to define and measure, and because its assumptions and predictions remain unclear. This is unfortunate because the RDH provides a potentially powerful model of grouping behavior in a diversity of conditions. Moreover, it can be generalized to predict other phenomena, including spacing behavior in nonsocial animals and utilization of resources other than food. Here, we review the empirical support, clarify the predictions of the RDH and argue that they can be used to provide better tests.     

Habitat selection and home range in the Blanford's fox,Vulpes cana: compatibility with the resource dispersion hypothesis

Summary This paper presents analyses of habitat-use and home range size in the Blanford's fox. We predicted, from the resource dispersion hypothesis (RDH), that home ranges would encompass similar areas of combined fruitful habitats, but widely different areas of useless habitats, and thus that home ranges would be larger where such fruitful patches are fragmented and widely dispersed. Home range estimates of 0.5–2.0 km² were calculated for 16 adult Blanford's foxes, using three different methods. There were no significant differences in home range size between sexes or study sites. One habitat, dry creekbed, was the most frequently visited in all home ranges. Dry creekbed provided abundant prey for the foxes and only sparse cover for their predators. Both the available area of creekbed in each range, and the area of creekbed patches that was used by the foxes, were independent of home range size. However, the variance in home range size was explained by the mean distance between the main denning area and the most frequently used patches of creekbed. These results are in accord with some predictions of the resource dispersion hypothesis.     

Movement patterns and herd dynamics among South African giraffes (Giraffa camelopardalis giraffa)

Giraffes reside in a fission–fusion social system, with sex, age proximity, kinship and home range overlap accounting for some of the variance in herd composition, while season, sex, age and time of day influence diet, home range size and distance travelled. To increase our knowledge of habitat use and fission–fusion herd dynamics, we placed GPS devices on eight adult female South African giraffes (Giraffa camelopardalis giraffa) living in the Khamab Kalahari Nature Reserve (South Africa). We tested four predictions about how season, kinship, home range and travel patterns influence habitat use and herd dynamics. Our two key findings were that females with a greater degree of home range overlap were more likely to form herds, but the degree of overlap was independent of the amount of time that they spent together in a herd, and that on the day prior to herd formation, females travelled about twice as far as their daily average and tended to move directly towards their future herd mate. We conclude that habitat use and movement patterns regulating fission–fusion dynamics reflect an interaction of ecological, social and reproductive factors operating in tandem, not independently.     

Only the largest terrestrial carnivores increase their dietary breadth with increasing prey richness

1. Animals should adapt their foraging habits, changing their dietary breadth in response to variation in the richness and availability of food resources. Understanding how species modify their dietary breadth according to variation in resource richness would support predictions of their responses to environmental changes that alter prey communities.
2. We evaluated relationships between the dietary breadth of large terrestrial carnivores and the local richness of large prey (defined as the number of species). We tested alternative predictions suggested by ecological and evolutionary theories: with increasing prey richness, species would (1) show a more diverse diet, thus broadening their dietary breadth, or (2) narrow their dietary breadth, indicating specialisation on a smaller number of prey.
3. We collated data from 505 studies of the diets of 12 species of large terrestrial mammalian carnivores to model relationships between two indices of dietary breadth and local prey richness.
4. For the majority of species, we found no evidence for narrowing dietary breadth (i.e. increased specialisation) with increasing prey richness. Although the snow leopard and the dhole appeared to use a lower number of large prey species with increasing prey richness, larger sample sizes are needed to support this result.
5. With increasing prey richness, the five largest carnivores (puma Puma concolor, spotted hyaena Crocuta crocuta, jaguar Panthera onca, lion Panthera leo, and tiger Panthera tigris), plus the Eurasian lynx Lynx lynx and the grey wolf Canis lupus (which are usually top predators in the areas from which data were obtained), showed greater dietary breadth and/or used a greater number of large prey species (i.e. increased generalism).
6. We suggest that dominant large carnivores encounter little competition in expanding their dietary breadth with increasing prey richness; conversely, the dietary niche of subordinate large carnivores is limited by competition with larger, dominant predators. We suggest that, over evolutionary time, resource partitioning is more important in shaping the dietary niche of smaller, inferior competitors than the niche of dominant ones.

     

Woodland fragmentation affects space use of Eurasian red squirrels

When habitats become fragmented, variation in patch size and quality are expected to impose changes on the spacing pattern and social organization of animal populations. General theory predicts different possible responses including shrinking home ranges (fission response), increasing range overlap (fusion) and incorporation of multiple patches in the home range (expansion response) as fragmentation increases. We studied space use and social organization in a metapopulation of red squirrels (Sciurus vulgaris) in 15 woodland fragments differing in size and tree species composition. Home ranges and core areas of males were larger than females, and fragmentation had different and complex effects on the spacing pattern of both sexes. In food-supplemented patches, high densities led to increased intra-sexual overlap. In linear-shaped patches, squirrels used smaller home ranges and core areas and had lower male–male and male–female overlap levels, independent of patch quality or size. Home range and core area size of males increased with patch size, and male core areas overlapped extensively those of other males and females. Hence males seemed to show a fission response only in some patches. In contrast, home range and core area size of females was not related with patch size, but decreased with habitat quality, supporting predictions of a fusion response and intra-sexual defense of food-based core areas. Hence, where patch size and shape strongly affected space use of male red squirrels, social organization of females was only affected in small, food-supplemented patches, suggesting that the basic spatio-social organization of adult females is very resistant to fragmentation.     

To eat and not be eaten: modelling resources and safety in multi-species animal groups

Using mixed-species bird flocks as an example, we model the payoffs for two types of species from participating in multi-species animal groups. Salliers feed on mobile prey, are good sentinels and do not affect prey capture rates of gleaners; gleaners feed on prey on substrates and can enhance the prey capture rate of salliers by flushing prey, but are poor sentinels. These functional types are known from various animal taxa that form multi-species associations. We model costs and benefits of joining groups for a wide range of group compositions under varying abundances of two types of prey-prey on substrates and mobile prey. Our model predicts that gleaners and salliers show a conflict of interest in multi-species groups, because gleaners benefit from increasing numbers of salliers in the group, whereas salliers benefit from increasing gleaner numbers. The model also predicts that the limits to size and variability in composition of multi-species groups are driven by the relative abundance of different types of prey, independent of predation pressure. Our model emphasises resources as a primary driver of temporal and spatial group dynamics, rather than reproductive activity or predation per se, which have hitherto been thought to explain patterns of multi-species group formation and cohesion. The qualitative predictions of the model are supported by empirical patterns from both terrestrial and marine multi-species groups, suggesting that similar mechanisms might underlie group dynamics in a range of taxa. The model also makes novel predictions about group dynamics that can be tested using variation across space and time.     

Effects of Individual and Group Characteristics on Feeding Behaviors in Wild Leontopithecus rosalia

Studies have linked variation in feeding and foraging success to variation in survival and reproductive success, which makes exploring influences on feeding invaluable. In the current study, we quantified energy contents of foods consumed by wild golden lion tamarins (Leontopithecus rosalia, GLT) and feeding behaviors of 34 GLT from March 1998 to March 1999. Our objective was to test predictions regarding effects of characteristics of the 1) individual, 2) group, 3) environment, and 4) other behaviors on 3 feeding behaviors: feeding on plant matter, searching for prey, and feeding on prey. We hypothesized that environmental characteristics, e.g., resource availability, in addition to group characteristics e.g., group size, would influence feeding on plant matter, because several individuals in a group often consume fruit in the same fruit tree. We hypothesized that environmental characteristics and individual characteristics, e.g., age, would influence searching for and consuming prey because the individual often searches for and consumes prey while it is alone at a substrate. We used SAS mixed models to determine the relative influence of these characteristics on the feeding behaviors. We found that group characteristics more significantly influenced feeding on plant matter, while individual characteristics more significantly influenced searching for prey. The results emphasize the distinctly different influences of individual and group characteristics on feeding. That influences other than competition may affect feeding on plant matter warrants further exploration.     

Size and scaling of predator-prey dynamics

We propose a scaled version of the Rosenzweig–MacArthur model using both Type I and Type II functional responses that incorporates the size dependence of interaction rates. Our aim is to link the energetic needs of organisms with the dynamics of interacting populations, for which survival is a result of a game-theoretic struggle for existence. We solve the scaled model of predator–prey dynamics and predict population level characteristics such as the scaling of coexistence size ranges and the optimal predator–prey size ratio. For a broad class of such models, the optimal predator–prey size ratio given available prey of a fixed size is constant. We also demonstrate how scaling predictions of prey density differ under resource limitation vs. predator drawdown. Finally, we show how evolution of predator size can destabilize population dynamics, compare scaling of predator–prey cycles to previous work, as well as discuss possible extensions of the model to multispecies communities.     

Dynamics of prey moving through a predator field: a model of migrating juvenile salmon

The migration of a patch of prey through a field of relatively stationary predators is a situation that occurs frequently in nature. Making quantitative predictions concerning such phenomena may be difficult, however, because factors such as the number of the prey in the patch, the spatial length and velocity of the patch, and the feeding rate and satiation of the predators all interact in a complex way. However, such problems are of great practical importance in many management situations; e.g., calculating the mortality of juvenile salmon (smolts) swimming down a river or reservoir containing many predators. Salmon smolts often move downstream in patches short compared with the length of the reservoir. To take into account the spatial dependence of the interaction, we used a spatially-explicit, individual-based modeling approach. We found that the mortality of prey depends strongly on the number of prey in the patch, the downstream velocity of prey in the patch, and the dispersion or spread of the patch in size through time. Some counterintuitive phenomena are predicted, such as predators downstream capturing more prey per predator than those upstream, even though the number of prey may be greatly depleted by the time the prey patch reaches the downstream predators. Individual-based models may be necessary for complex spatial situations, such as salmonid migration, where processes such as schooling occur at fine scales and affect system predictions. We compare some results to predictions from other salmonid models.     

Opportunistic feeding by lions: non-preferred prey comprise an important part of lion diets in a habitat where preferred prey are abundant

Optimal foraging theory predicts less diverse predator diets with a greater availability of preferred prey. This narrow diet niche should then be dominated by preferred prey, with implications for predator–prey dynamics and prey population ecology. We investigated lion (Panthera leo) diets in Hluhluwe–iMfolozi Park (HiP), South Africa, to assess whether lions in a site with a high density of preferred prey (prey species weighing 92–632 kg as estimated from a published meta-analysis) have a narrow diet, consisting primarily of preferred prey. HiP is a useful study site to investigate this prediction because it is a productive landscape (with a high density of prey) where lion-preferred prey constitutes up to 33% of the prey available to lions. Furthermore, to investigate whether lions in HiP exhibit sex-specific diets as documented in other southern African populations, we estimated male and female lion diets separately. We were specifically interested in testing whether traditional approaches of estimating lion diets at the population level mask sex-specific predation patterns, with possible implications for management of lions in small to medium-sized fenced reserves. Lions in HiP preferred larger prey species (63–684 kg) and had diets with a larger proportion of preferred prey than reported in an African-wide meta-analysis. However, despite the high density of preferred prey species, 36% of lion diets still consisted of typically non-preferred species such as nyala (Tragelaphus angasii). This finding suggests that lions in HiP maintain a degree of opportunism even when preferred prey are abundant. Therefore, abundant, non-preferred prey are likely to be an important resource for lion populations. Sex-specific differences in lion diets were evident in HiP, suggesting that estimation of lion resource use and carrying capacity should consider opportunistic hunting and sex-specific differences in lion diets.     

Spatial heterogeneity of biomass turnover has contrasting effects on synchrony and stability in trophic metacommunities

Spatial heterogeneity is a fundamental feature of ecosystems, and ecologists have identified it as a factor promoting the stability of population dynamics. In particular, differences in interaction strengths and resource supply between patches generate an asymmetry of biomass turnover with a fast and a slow patch coupled by a mobile predator. Here, we demonstrate that asymmetry leads to opposite stability patterns in metacommunities receiving localized perturbations depending on the characteristics of the perturbed patch. Perturbing prey in the fast patch synchronizes the dynamics of prey biomass between the two patches and destabilizes predator dynamics by increasing the predator's temporal variability. Conversely, perturbing prey in the slow patch decreases the synchrony of the prey's dynamics and stabilizes predator dynamics. Our results have implications for conservation ecology and suggest reinforcing protection policies in fast patches to dampen the effects of perturbations and promote the stability of population dynamics at the regional scale.     

Effects of Individual and Group Characteristics on Feeding Behaviors in Wild <Emphasis Type="Italic">Leontopithecus rosalia</Emphasis>

Studies have linked variation in feeding and foraging success to variation in survival and reproductive success, which makes exploring influences on feeding invaluable. In the current study, we quantified energy contents of foods consumed by wild golden lion tamarins (Leontopithecus rosalia, GLT) and feeding behaviors of 34 GLT from March 1998 to March 1999. Our objective was to test predictions regarding effects of characteristics of the 1) individual, 2) group, 3) environment, and 4) other behaviors on 3 feeding behaviors: feeding on plant matter, searching for prey, and feeding on prey. We hypothesized that environmental characteristics, e.g., resource availability, in addition to group characteristics e.g., group size, would influence feeding on plant matter, because several individuals in a group often consume fruit in the same fruit tree. We hypothesized that environmental characteristics and individual characteristics, e.g., age, would influence searching for and consuming prey because the individual often searches for and consumes prey while it is alone at a substrate. We used SAS mixed models to determine the relative influence of these characteristics on the feeding behaviors. We found that group characteristics more significantly influenced feeding on plant matter, while individual characteristics more significantly influenced searching for prey. The results emphasize the distinctly different influences of individual and group characteristics on feeding. That influences other than competition may affect feeding on plant matter warrants further exploration. An erratum to this article can be found at     

Mechanisms of coexistence in diverse herbivore–carnivore assemblages: demographic,temporal and spatial heterogeneities affecting prey vulnerability

Simple models coupling the dynamics of single predators to single prey populations tend to generate oscillatory dynamics of both predator and prey, or extirpation of the prey followed by that of the predator. In reality, such oscillatory dynamics may be counteracted by prey refugia or by opportunities for prey switching by the predator in multi‐prey assemblages. How these mechanisms operate depends on relative prey vulnerability, a factor ignored in simple interactive models. I outline how compositional, temporal, demographic and spatial heterogeneities help explain the contrasting effects of top predators on large herbivore abundance and population dynamics in species‐rich African savanna ecosystems compared with less species‐diverse northern temperate or subarctic ecosystems. Demographically, mortality inflicted by predation depends on the relative size and life history stage of the prey. Because all animals eventually die and are consumed by various carnivores, the additive component of the mortality inflicted is somewhat less than the predation rate. Prey vulnerability varies annually and seasonally, and between day and night. Spatial variation in the risk of predation depends on vegetation cover as well as on the availability of food resources. During times of food shortage, herbivores become prompted to occupy more risky habitats retaining more food. Predator concentrations dependent on the abundance of primary prey species may restrict the occurrence of other potential prey species less resistant to predation. The presence of multiple herbivore species of similar size in African savannas allows the top predator, the lion, to shift its prey selection flexibly dependent on changing prey vulnerability. Hence top–down and bottom–up influences on herbivore populations are intrinsically entangled. Models coupling the population dynamics of predators and prey need to accommodate the changing influences of prey demography, temporal variation in environmental conditions, and spatial variation in the relative vulnerability of alternative prey species to predation. Synthesis While re‐established predators have had major impacts on prey populations in northern temperate regions, multiple large herbivore species typically coexist along with diverse carnivores in African savanna ecosystems. In order to explain these contrasting outcomes, certain functional heterogeneities must be recognised, including relative vulnerability of alternative prey, temporal variation in the risk of predation, demographic differences in susceptibility to predation, and spatial contrasts in exposure to predation. Food shortfalls prompt herbivores to exploit more risky habitats, meaning that top–down and bottom–up influences on prey populations are intrinsically entangled. Models coupling the interactive dynamics of predator and prey populations need to incorporate these varying influences on relative prey vulnerability.