Ranging behavior of the mountain gorillas (Gorilla beringei beringei) in Bwindi Impenetrable National Park, Uganda: a test of the ecological constraints model

The ecological constraints model predicts that daily travel distance and home range size of social animals will increase as group size increases in order to meet the dietary needs of additional group members. This theory has been supported more predominantly by studies of frugivorous primate species than by studies of folivorous species. We examined the ranging patterns of mountain gorillas in Bwindi Impenetrable National Park, Uganda, who include both herbaceous vegetation and fruit in their diet, to determine how ecological, behavioral, and social parameters influence movement patterns. Data were collected from three groups of gorillas with overlapping home ranges at a low-altitude location (1,450–1,800 m) and one group at a high-altitude location (2,100–2,500 m) in Bwindi from September 2001 to August 2002. We analyzed daily travel distance and home range size in relation to group size, while also considering patterns of frugivory, rainfall, and location (proxy for food availability) within the park. Both daily travel distance and home range size were positively related to group size. In addition, the degree of frugivory positively influenced daily travel distance and home range size, while rainfall negatively influenced daily travel distance only. Finally, groups at the low-altitude location, with higher fruit availability, traveled less than the group at the high-altitude location. These results demonstrate that mountain gorillas in Bwindi provide support for the ecological constraints model, but further studies are needed to determine how fine-scale spatial and temporal availability of food resources influence movement patterns. Ranging patterns of Bwindi gorillas are compared to those observed in other gorilla populations in the context of the ecological constraints model.Communicated by D. Watts     

Modeling the Persistence of Small Populations of Strongly Interdependent Species: Figs and Fig Wasps

Conservation problems are usually studied at the population or ecosystem levels. Formulating predictive theory for the domain in between has been difficult. Fig trees and their pollinating wasps, principally tropical groups of organisms, form pairs of obligate mutualists that provide unique opportunities for studying the influence of species interactions on the survival of small populations. Survival of each partner depends on that of the associated species. The pollinator population can be maintained only if figs are produced year-round. Because fig trees flower synchronously at the individual level, wasps have to locate a new individual host tree at each generation. We describe results of simulation models estimating the minimum number of trees required to maintain a wasp population using two levels of the criteria: (1) different probability of survival (50% and 99%) and (2) different time of survival (5 or 1000 years). We also examined how these different estimates are sensitive to differences in the seasonality of flowering period and in the length of the period of female receptivity in figs. Such estimates can be used to understand the potential effects of the reduction of fig population size via fragmentation. Unlike most studies on the effect of low population size on population viability, our paper focuses on maintenance of a biotic interaction, rather than on single-species dynamics. The biotic interaction on which we focus is important because figs in many tropical ecosystems may be keystone resources for frugivores that are in turn essential seed dispersal agents for other plants.     

Female philopatry and its social benefits among Bornean orangutans

Female philopatry in mammals is generally associated with ecological and sometimes social benefits, and often with dispersal by males. Previous studies on dispersal patterns of orangutans, largely non-gregarious Asian great apes, have yielded conflicting results. Based on 7?years of observational data and mitochondrial and nuclear DNA analyses on fecal samples of 41 adult Bornean orangutans (Pongo pygmaeus wurmbii) from the Tuanan population, we provide both genetic and behavioral evidence for male dispersal and female philopatry. Although maternally related adult female dyads showed similar home-range overlap as unrelated dyads, females spent much more time in association with known maternal relatives than with other females. While in association, offspring of maternally related females frequently engaged in social play, whereas mothers actively prevented this during encounters with unrelated mothers, suggesting that unrelated females may pose a threat to infants. Having trustworthy neighbors may therefore be a social benefit of philopatry that may be common among solitary mammals, thus reinforcing female philopatric tendencies in such species. The results also illustrate the diversity in dispersal patterns found within the great-ape lineage.     

The development of sex differences in ring-tailed lemur feeding ecology

Sex differences in feeding ecology may develop in response to fluctuations in physiological costs to females over their reproductive cycles, or to sexual size dimorphism, or function to minimize feeding competition within a group via resource partitioning. For most mammal species, it is unknown how these factors contribute to sex differences in feeding, or how the development of males and females reflects these intraspecific feeding differences. We show changes in dietary composition, diversity, overlap, and foraging behavior throughout development in ring-tailed lemurs (Lemur catta) and test how the development of sex differences in feeding is related to female costs of reproduction and year-round resource partitioning. Sex differences in dietary composition were only present when females were lactating, but sex differences in other aspects of feeding, including dietary diversity, and relative time spent feeding and foraging, developed at or near the time of weaning. Sex difference in juveniles and subadults, when present, were similar to the differences found in adults. The low year-round dietary overlap and early differences in dietary diversity indicate that some resource partitioning may begin with young individuals and fluctuate throughout development. The major differences between males and females in dietary composition suggest that these larger changes in diet are closely tied to female reproductive state when females must shift their diet to meet energetic and nutritional requirements.     

On the possible contribution of mixed species flocks to species richness in neotropical avifaunas

Summary Mixed species foraging flocks are a dominant component of the infra-structure of avian communities in neotropical forests. In Amazonia, these flocks consist of pairs of 10–20 species, many of which are permanently associated with mixed flocks. At least half of these flocking species maintain territories that correspond exactly to the flock home range. Small individuals that participate as permanent members of the flocks must adopt the large home range of the larger nucleus species. Therefore, the densities of smaller species are dependent on the availability and density of flocks rather than the availability of food resources. Single pairs of 4 small flocking species with individual body masses of 8 g occupied exclusive territories of 8–12 ha. These were the same exact territories that were defended by at least 6 other flocking species with individual body masses of up to 37 g. Because of their attachment to flocks with large territories, small species are expected to under-utilize available food resources. The under-utilization of food resources is expected to allow smaller species to coexist with greater niche overlap resulting in increased species richness. This hypothesis was tested by quantifying foraging niche in terms of foraging height, foraging maneuver, and prey substrate; and using these values in addition to body mass and bill size (length, depth and width) to determine relative niche overlap between large versus small species pairs.Smaller species had greater foraging overlap than large flocking species and particularly the three smallest species of the genus Myrmotherula; longipennis, axillaris and menetriesii had very high overlap (average foraging niche overlap for the 3 species=0.83±0.12 compared with 0.12±0.19 for all flocking species), similar body sizes (body masses differing by no more then 8%) and similar bill morphologies (maximum ratio in length=1.08, width=1.07, and depth=1.06). These results are consistent with the hypothesis that small species participating in Amazonian mixed flocks can coexist with greater niche overlap because their density is flock dependent rather than resource dependent.     

Use of heterospecific cues by the lizard Platysaurus broadleyi for food location

Flat lizards (Platysaurus broadleyi) at Augrabies Falls National Park, South Africa, are restricted to rocky terrain where the predominant tree is the Namaqua fig (Ficus cordata cordata). P. broadleyi readily feeds on Namaqua figs when they are available, and the lizards sometimes form large (maximum recorded=134) congregations under fruiting fig trees. The distance lizards travel to fruiting trees also exceed normal daily foraging distances. Location of fruiting fig trees by a lizard can have a high pay-off because figs are energetically rich and trees fruit irregularly and asynchronously, resulting in a resource that is available, but unpredictable in time and space. The prediction that bird activity in fig trees provides a cue to the presence of ripe figs was tested experimentally. By placing cages containing birds and empty control cages in trees devoid of fruit, we demonstrated that P. broadleyi are drawn to fig trees with high bird congregations. We also tested if the presence of a fig tree was necessary to draw lizards to bird congregations by placing cages containing birds and empty control cages in a matched-pairs design on rock away from trees. Namaqua fig trees were not necessary to draw lizards to bird congregations. Received: 12 May 1998 / Accepted after revision: 30 November 1998     

Feeding ecology of three species of Astropecten (Asteroidea) coexisting on shallow sandy bottoms of the northwestern Mediterranean Sea

Predation and competition are important biotic interactions influencing populations and communities in marine soft sediments. Sea stars are ubiquitous predators with diverse diets that play functionally important roles in the benthos. In this study, we examined the diet and the ecological roles of three sympatric species of the genus Astropecten (A. aranciacus, A. irregularis pentacanthus and A. platyacanthus). The study was performed between March 2010 and February 2011 on the Maresme coast (northwestern Mediterranean Sea). Results showed that their main diet consisted on gastropods and bivalves, such as Glycymeris glycymeris, Callista chione, Gibbula guttadauri and Cyclope neritea. Food competition between species was avoided by partition of prey resources. Intraspecific differences in the dietary compositions between seasons were found, but not between size classes. Ontogenetic patterns of prey size consumption were recognized in the three species. A large diet overlap was detected between A. aranciacus and A. platyacanthus in winter, due to changes in prey availability. Nevertheless, the analysis of the infaunal community composition and stomach contents indicated that food selection was not associated with prey availability.     

Determinants of male spacing behavior in microtines: an experimental manipulation of female spatial distribution and density

Theory on microtine mating systems predicts that male spacing behavior will be related to female spatial and temporal distribution. However, data from a natural population of field voles, Microtus agrestis, indicated a potential influence of female density on the spacing behavior of males. Therefore, I experimentally investigated the relative importance of female density and female spatial distribution for the spacing behavior of males in M. agrestis. Males were radio-tracked in enclosed natural habitats in which females at different densities were placed in two different spatial arrangements: clumped versus even distribution. Female density was the main factor determining male spacing behavior. At the high female density males had smaller home ranges and moved shorter distances between radio-tracking recordings. Also, home ranges were more exclusive at high female density. However, since there was a significant positive correlation between home range size and range overlap, range exclusiveness seemed to be influenced by female density indirectly through the effect of range size. Female spatial distribution, on the other hand, had no influence on male home range exclusiveness.     

Reproductive trade-offs from mating with a successful male: the case of the tephritid fly Anastrepha obliqua

In lekking species, females may become sperm-limited when mating with sexually successful males, and this may be exacerbated by a poor male diet. Polygynous males may also be limited by the amount of accessory gland products (AGPs) they can transmit to females, which in turn may influence the females’ refractory period and longevity. Here, we tested the effect of male mating history, larval and adult diet on copula duration, mating intervals, female fecundity, fertilisation success, life span and likelihood to remate using sexually successful males of the lekking tephritid fly Anastrepha obliqua. Flies originated from either a native or exotic host fruit and were protein-fed or deprived. Male diet and larval host influenced copula duration, while the time elapsed between matings was affected by the interaction of mating order and male adult diet. Female fecundity was not influenced by female position in mating order or protein inclusion into the male diet. However, mating order and male larval diet influenced female fertilisation success. Importantly, as males mated successively they were less able to induce a refractory period on females, as the last females to mate with a male were more likely to remate and had slightly longer life spans than the first females to mate with males. These results might be attributed to a decrease in male AGPs with increasing male mating frequency. We discuss the role of conditional expression of male mating frequency with respect to A. obliqua’s life history, the trade-off that females face when mating with a successful male, the effect of larval diet on adult sexual performance and the possibility for sexual conflict to occur due to high male mating rates and fitness costs to females.     

Geographical gradients in diet affect population dynamics of Canada lynx

Geographical gradients in the stability of cyclic populations of herbivores and their predators may relate to the degree of specialization of predators. However, such changes are usually associated with transition from specialist to generalist predator species, rather than from geographical variation in dietary breadth of specialist predators. Canada lynx (Lynx canadensis) and snowshoe hare (Lepus americanus) populations undergo cyclic fluctuations in northern parts of their range, but cycles are either greatly attenuated or lost altogether in the southern boreal forest where prey diversity is higher. We tested the influence of prey specialization on population cycles by measuring the stable carbon and nitrogen isotope ratios in lynx and their prey, estimating the contribution of hares to lynx diet across their range, and correlating this degree of specialization to the strength of their population cycles. Hares dominated the lynx diet across their range, but specialization on hares decreased in southern and western populations. The degree of specialization correlated with cyclic signal strength indicated by spectral analysis of lynx harvest data, but overall variability of lynx harvest (the standard deviation of natural-log-transformed harvest numbers) did not change significantly with dietary specialization. Thus, as alternative prey became more important in the lynx diet, the fluctuations became decoupled from a regular cycle but did not become less variable. Our results support the hypothesis that alternative prey decrease population cycle regularity but emphasize that such changes may be driven by dietary shifts among dominant specialist predators rather than exclusively through changes in the predator community.     

Multi-year lactation and its consequences in Bornean orangutans (Pongo pygmaeus wurmbii)

In most mammals, females pay for reproduction by dramatically increasing net energy intake from conception to mid- or late lactation. To do this, they time their reproductive events in relation to environmental cycles so that periods of peak food availability coincide with peak demand or are used to build energy stores. This timing is not possible in species with slow development in which lactation is prolonged over a multi-year period with fluctuating food availability. Here, mothers are expected to sustain a stable but generally lower level of nutrient transfer. In a sample of over 1,050 complete follow days of eight mother–infant pairs collected over 7 years, we document maternal effort for wild Bornean orangutans (Pongo pygmaeus wurmbii) over their average 6.5-year lactation period. As predicted, maternal feeding time was independent of the age of her growing offspring, indicating a stable sustained “plateau” effort of ≤ 25 % above baseline level, instead of a short peak lactation as seen in seasonal breeders. Infant orangutans started to regularly supplement milk with self-harvested food when they were 1–1.5 years old, indicating milk intake was insufficient from this age onwards, even though maternal effort did not decrease. We expect the same regulation of sustained maternal effort in other large and large-brained mammals with slow infant development. We also predict that mother–infant conflict over suckling may show another peak at the onset of the milk?+?solid food phase, in addition to the well-known conflict around the endpoint of lactation (weaning), which is reached after a long and gradual increase in solid food intake by the infant.     

Female tactics to reduce sexual harassment in the Sumatran orangutan (Pongo pygmaeus abelii)

Orangutan males demonstrate intrasexual dimorphism with corresponding alternative mating strategies. Sexual harassment is the predominant feature of the mating strategy that subadult males pursue. This study investigated the countertactics that females employ to reduce sexual harassment by subadult males. I observed 207 copulations during more than 9,000 h of focal follows of wild Sumatran orangutans at the Suaq Balimbing Research Station over a 23-month period. Rates of copulations initiated by subadult males increased during months of high fruit abundance, and most mating attempts were directed toward females with weaned infants. Simultaneous harassment by multiple subadult males increased significantly during months of high fruit abundance, and nearly all adult female-adult male consortships occurred during periods of high fruit abundance. Females who maintained spatial association with adult males, either via consortship or by nonmating temporary parties, received lower rates of harassment, as measured by the success rate of subadult male mating attempts. Adult female-adult male parties did not always result in mating between the associating dyad. Female initiation of protective services by adult males is one social tactic that female orangutans employ to reduce sexual harassment. Females therefore receive direct services from adult males, which may be one factor that influences female mate choice in Sumatran orangutans.     

The trophic ecology of two abundant mesopredators in south-east coastal waters of Tasmania, Australia

Investigating predator–prey relationships is an important component for identifying and understanding the factors that influence the structure and function of ecosystems. Mesopredators, defined as mid-level predators, have a profound effect on ecosystem structure by contributing an important link between apex predators and lower trophic levels. The diet of two elasmobranch mesopredators, Squalus acanthias and Mustelus antarcticus, was investigated in three locations in south-east Tasmania. Squalus acanthias consumed predominantly pelagic teleosts and cephalopods, while M. antarcticus predominantly consumed benthic crustaceans. As a result, there was low dietary and niche overlap between the two species. There was however evidence of intra-specific dietary variations between locations for both the species. This study has contributed to a better understanding of the top-down dynamics of the food web in coastal Tasmania, by providing important dietary information of two abundant mesopredators. In addition, the similar dietary patterns for S. acanthias and other Mustelus species over much of their global range suggest they may be consistent in their trophic roles across systems, with limited competition between these two sympatric mesopredators to be expected.     

Urbanized landscapes favored by fig-eating birds increase invasive but not native juvenile strangler fig abundance

Propagule pressure can determine the success or failure of invasive plant range expansion. Range expansion takes place at large spatial scales, often encompassing many types of land cover, yet the effect of landscape context on propagule pressure remains largely unknown. Many studies have reported a positive correlation between invasive plant abundance and human land use; increased propagule pressure in these landscapes may be responsible for this correlation. We tested the hypothesis that increased rates of seed dispersal by fig-eating birds, which are more common in urban habitats, result in an increase in invasive strangler fig abundance in landscapes dominated by human land use. We quantified abundance of an invasive species (Ficus microcarpa) and a native species (F. aurea) of strangler fig in plots spanning the entire range of human land use in South Florida, USA, from urban parking lots to native forest. We then compared models that predicted juvenile fig abundance based on distance to adult fig seed sources and fig-eating bird habitat quality with models that lacked one or both of these terms. The best model for juvenile invasive fig abundance included both distance to adult and fig-eating bird habitat terms, suggesting that landscape effects on invasive fig abundance are mediated by seed-dispersing birds. In contrast, the best model for juvenile native fig abundance included only presence/absence of adults, suggesting that distance from individual adult trees may have less effect on seed limitation for a native species compared to an invasive species undergoing range expansion. However, models for both species included significant effects of adult seed sources, implying that juvenile abundance is limited by seed arrival. This result was corroborated by a seed addition experiment that indicated that both native and invasive strangler figs were strongly seed limited. Understanding how landscape context affects the mechanisms of plant invasion may lead to better management techniques. Our results suggest that prioritizing removal of adult trees in sites with high fig-eating bird habitat may be the most effective method to control F. microcarpa abundance.     

Dynamics of biparental care in the dampwood termite,Zootermopsis nevadensis (Hagen): response to nitrogen availability

Summary Studies were conducted on the dampwood termite, Zootermopsis nevadensis, to examine the behavioral roles of the reproductive pair during the nest-founding period and to determine the effect of nitrogen availability on their reproduction and division of labor. Nitrogen has been hypothesized to be an important limiting nutrient for founding pairs. One nitrogenous reserve, uric acid-nitrogen, was examined in reproductives and in nutrient-receiving and nutrient-gathering colony members; it was found in highest amounts in the reproductives (i.e., alates, de-alates, and primary reproductives). Young pairs may use these nitrogenous reserves to increase their chances for reproduction. In support of this hypothesis, founding pairs that were fed a diet supplemented with uric acid-nitrogen had a significantly greater probability of producing at least one offspring than did pairs fed an unsupplemented diet. Females that were fed a nitrogen-poor diet restricted their total activity while their mates sustained a high activity during colony initiation. When fed a nitrogen-rich diet, females collected pulp more often than their mates, while males collected more water, though only in the period prior to egg laying. In all pairs, males transferred proctodeal pellets (food derived from the hindgut intestine) to their mates significantly more often than females to males, and females fed on proctodeal pellets significantly more often than did their mates. The male-female asymmetries in pellet transfer and feeding were not significant in the stage after egg deposition. Once eggs and larvae were present in the nest, a male and female spend an equal percentage of time caring for eggs and feeding larvae. Proctodeal pellets examined in reproductives were found to be rich in proteins. This suggests that in the pre-egg period, the male provides nitrogen-rich substances to the female as a form of paternal investment.     

Growing of trees in home-gardens by rural households in the Eastern Cape and Limpopo Provinces,South Africa

Trees provide a wide range of goods and services to rural households which, when incorporated into their livelihood strategies, help reduce their vulnerability to adversity. Governments and policy makers often ignore the contribution made by trees and consequently resources are focussed on cash crops and livestock. Villagers in the Eastern Cape and Limpopo Province, South Africa utilise a range of trees from home-gardens for various purposes, although predominantly for fruit and shade. Trees are either planted or actively retained in households' home-gardens. There were noticeable differences between the villages in the Eastern Cape and those in Limpopo Province, particularly with respect to the overall density of trees per hectare and the number of species per household, both being significantly greater in Limpopo Province. The five most preferred species were listed for each village, revealing a preference for exotic fruit trees in Limpopo Province and a mix of exotic fruit trees and shade trees in the Eastern Cape. Households also retained useful indigenous species, predominantly fruit-bearing species. A range of factors constrain tree growing in home-gardens and households engage in practices to grow and maintain their trees. Not all of these constraints and practices were significantly different between the various localities.     

The nature and frequency of social interactions among free-living prairie voles (Microtus ochrogaster)

Hypotheses relating the behavior of voles to their population cycles often assume that the rate of social interaction increases with population density. To test this assumption, we examined the frequency of social interactions in a population of prairie voles (Microtus ochrogaster) over a 7-year period. In addition, we characterized space use by resident animals, patterns of visitation by nonresidents to nests, and participants in social interactions. Social groups within the population typically displayed little overlap in their use of space, even at high population densities. Nevertheless, nonresidents, particularly wandering males, were captured as visitors at nests. The number of visits per social group did not increase in a simple linear manner with population density and was particularly variable when there were fewer than 100 animals/ha. At such times, more single females and fewer pairs received visits from males than expected based on the frequency of occurrence of these groups in the population; a similar pattern was noted during periods of high population density (≥100 animals/ha) but the comparisons failed to reach statistical significance. Furthermore, at high population density, more communal groups received visits from females than expected. Patterns of visitation to communal groups were influenced by the number of adult male residents (winter only), but not by the number of adult female residents or presence of philopatric female offspring. These data indicate that the frequency of social contact in prairie voles does not increase linearly with population density and is influenced by the spacing and possible mate-guarding behavior of resident animals. Received: 7 January 1998 / Accepted after revision: 16 May 1998     

A Landscape Approach to Biodiversity Conservation of Sacred Groves in the Western Ghats of India

Abstract:  Although sacred groves are important for conservation in India, the landscape that surrounds them has a vital influence on biodiversity within them. Research has focused on tree diversity inside these forest patches. In a coffee-growing region of the Western Ghats, however, landscape outside sacred groves is also tree covered because planters have retained native trees to provide shade for coffee plants. We examined the diversity of trees, birds, and macrofungi at 58 sites—10 forest-reserve sites, 25 sacred groves, and 23 coffee plantations— in Kodagu district. We measured landscape composition and configuration around each site with a geographic information system. To identify factors associated with diversity we constructed multivariate models by using a decision-tree technique. The conventional measures of landscape fragmentation such as patch size did not influence species richness. Distance of sacred groves from the forest reserve had a weak influence. The measures of landscape structure (e.g., tree cover in the surroundings) and stand structure (e.g., variability in canopy height) contributed to the variation in species richness explained by multivariate models. We suggest that biodiversity present within sacred groves has been influenced by native tree cover in the surrounding landscape. To conserve this biodiversity the integrity of the tree-covered landscape matrix will need to be conserved.     

Feeding opportunities of larval and juvenile cod (Gadus morhua) in a Greenlandic fjord: temporal and spatial linkages between cod and their preferred prey

Feeding of fish depends on a spatial and temporal match with prey, and since larval and juvenile feeding can be highly selective, their preferences for given prey sizes and taxa should be considered when quantifying the actual availability of potential prey. We investigated the diet and prey preferences of the early-life stages of Atlantic cod (Gadus morhua) to quantify the availability of prey during a spring-summer season in a West Greenlandic fjord. We hypothesized that abundances of larval and juvenile cod at size were synchronized to optimal availability of preferred prey in space and time. The present analysis is based on nine cruises each covering 5 stations visited between 24 May and 5 August 2010 comparing zooplankton abundance, cod gut content and distribution patterns. Cod 4–25 mm in length preferred prey of about 5 % of their own length. During ontogeny, their preferences changed from calanoid nauplii towards Pseudocalanus spp. and Calanus spp. copepodites. The larvae/juvenile had an exceptionally high dietary contribution from cladocerans, which were highly preferred by cod larger than 9 mm, while the abundant Metridia longa and the non-calanoid copepods contributed less. These findings stress the importance of focusing on abundance of preferred prey when assessing the actual prey availability to young fish. We found a spatio-temporal overlap between cod and their preferred prey, and observations suggest that advection of both zooplankton and cod contributed to this overlap. Hence, the larval feeding opportunities might be sensitive to climate-related changes affecting the circulation patterns in this fjord.     

Density dependence in social behaviour: home range overlap and density interacts to affect conspecific encounter rates in a gregarious ungulate

Sociality is poorly understood in the context of population processes. We used wild, female elk (Cervus canadensis) equipped with proximity-logging radio collars (n?=?62) from Manitoba, Canada (2007–2009), to test for modifying effects of population density (two areas: 0.42 and 0.22 animals/km²) on the relationship between two measures of sociality. This included the rate at which collared individuals encountered one another per year (encounters logged as animals ranging to within 1.4 m of each other) and the extent to which animals overlapped in annual home range (proportion of shared minimum convex polygon ranges). Overlap was significantly greater in the high density area compared to that of the low, but not if we only considered individuals that directly encountered each other, implying that familiar individuals will maintain a constant degree of range overlap regardless of density. Encounter rate was nonlinearly related to home range overlap. This relationship was also density-dependent, exhibiting negative density dependence at high proportions of overlap, primarily in the high density subpopulation. Sociality, as defined by two interacting measures of behaviour—encounter rate and home range overlap—exhibits a complex nonlinear relationship; we discuss the implications of these results as they pertain to sociobiology, resource competition, and pathogen transmission.