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A B S T R A C T The use of social network theory in evaluating animal social groups has gained traction in recent years. Despite the utility of social network analysis in describing attributes of social groups, it remains unclear how comparable this approach is to traditional behavioral observational studies. Using data on Gunnison's prairie dog (Cynomys gunnisoni) social interactions we describe social networks from three populations. We then compare those social networks to groups identified by traditional behavioral approaches and explore whether individuals group together based on similarities. The social groups identified by social network analysis were consistent with those identified by more traditional behavioral approaches. However, fine-grained social sub-structuring was revealed only with social network analysis. We found variation in the patterns of interactions among prairie dog social groups that was largely independent of the behavioral attributes or genetics of the individuals within those groups. We detected that some social groups include disproportionately well-connected individuals acting as hubs or bridges. This study contributes to a growing body of evidence that social networks analysis is a

Our species displays remarkable linguistic diversity. Although the uneven distribution of this diversity demands explanation, the drivers of these patterns have not been conclusively determined. We address this issue in two steps: First, we review previous empirical studies whose authors have suggested environmental, geographical, and sociocultural drivers of linguistic diversification. However, contradictory results and methodological variation make it difficult to draw general conclusions. Second, we outline a program for future research. We suggest that future analyses should account for interactions among causal factors, the lack of spatial and phylogenetic independence of the data, and transitory patterns. Recent analytical advances in biogeography and evolutionary biology, such as simulation modeling of diversity patterns, hold promise for testing four key mechanisms of language diversification proposed here: neutral change, population movement, contact, and selection. Future modeling approaches should also evaluate how the outcomes of these processes are influenced by demography, environmental heterogeneity, and time.

Sexually selected traits and the use of strategies to enhance male reproductive success (e.g., competition and dispersal) can yield sex differences in metabolic requirements, rates and durations of growth and maturation, and the propensity for risky behavior, which are suggested to result in age-specific sex differences in mortality and life span. We investigated age-specific sex ratios, mortality, and dispersal in Propithecus edwardsi in Ranomafana National Park, Madagascar. We predicted that, due to similarities in growth rates and body sizes, male and female juvenile mortality rates would be comparable; because both sexes disperse and have intense intersexual competition and aggression, adult mortality would be similar; given similarities in dispersal frequency and distance, the timing of dispersal would not differ. We used 80 group-years births, deaths, and dispersals (N females = 41, N males = 34) collected over 23 years to calculate sex ratios and survival curves. Females lived longer than males (maximum 32 and 19 years, respectively). Sex ratios were male biased from sexual maturity through 17 years and female biased at birth and older ages. Infant survival probabilities were similar. Thus, differential development and maturation are unlikely explanations for longer female life span in this species. Males were more likely to survive from 2 to 18 years. However, male annual survival probability declined quickly around 13–18 years; males continued to disperse until their deaths, whereas females generally stopped dispersing after 11 years. We suggest that sex differences in the timing of dispersal and the unique challenges of risky behavior at older ages may be sufficient to yield differences in male and female life span.

In many social sciurids, male territoriality confers significant mating advantages. We evaluated resident male paternity in Gunnison's prairie dogs (Cynomys gunnisoni), a colonial ground-dwelling sciurid, where males and females cooperatively defend territories. Contrary to findings reported for other social sciurids, our results show that territorial resident males do not gain significant reproductive advantages. Resident males sired the majority of offspring from their respective territories only 10.5% of the time. A single non-resident male sired equal or greater number of offspring than any single resident male 71.2% of the time. While adult males were more likely to sire a greater number of offspring, standard heterozygosity, body mass, and territory size were not significant predictors of how many offspring a male sired. In addition, the number of other males present did not influence the number of offspring sired by any given resident male. However, territory size was significantly correlated with overall offspring number and mean litter size per female, suggesting a potential reproductive advantage for females that occupy larger, better quality territories. Previous work has demonstrated that the cooperative defence of territories by both males and females permits enhanced access to food resources critical to overwinter survival. Our results have important implications for studies that do not distinguish between social and mating systems, because they suggest that individual mating strategies may be different from the social strategies that emerge in response to resource availability.

Few studies have experimentally tested the resource dispersion hypothesis (RDH). In this study, I tested whether space use and social organization of Gunnison’s prairie dog responded to changes in the dispersion and abundance of resources. Food manipulations were carried out during the reproductive and nonreproductive seasons across 2 years. Gunnison’s prairie dog adults responded to the experiments by decreasing territory size as food became patchier in space and time. Both males and females modified their home ranges, with no detectable difference between sexes, either prior to or during the experiments. As food became patchier in space and time, the spatial overlap of adults increased, whereas it decreased as food became more evenly dispersed. The average size of a group, defined as those individuals occupying the same territory, did not change significantly as a result of the experiments. Where changes in the composition and size of groups did occur, there was no indication that such changes were sex specific. Results from this study support critical components of the RDH and strongly suggest that patterns of space use and social structure in Gunnison’s prairie dogs are the result of individual responses to resource abundance and distribution.

In this study, we describe patterns of relatedness in Gunnison’s prairie dog (Cynomys gunnisoni) social groups. Kin selection is often cited as a mechanism for the evolution and maintenance of social groups, and Gunnison’s prairie dog females are occasionally described as being strongly philopatric. Overall, randomization tests revealed that females within territorial groups were not more closely related to each other than expected at random. A similar pattern was found among males and between males and females, indicating that there was no sex-biased dispersal occurring in these populations. Ecological variables measured in this study, such as food abundance and food dispersion, were not correlated with relatedness. In addition, territory size and density ⁄m 2 did not correlate significantly with relatedness. Although there was variability in the spatial overlap among individuals within groups, there was no indication that relatedness explained this variation. These results suggest that kin selection is not maintaining social groups in these populations, but that competition for access to resources required by both males and females may explain dispersal and social group patterns in these populations.

In this paper, we provide a description of six excavated burrow systems of Gunnison's prairie dog (Cynomys gunnisoni) in northern Arizona. These data plus other data from the literature on structure of the burrow system of Gunnison's prairie dogs, black-tailed prairie dogs (C. ludovicianus), white-tailed prairie dogs (C. leucurus), and Utah prairie dogs (C. parvidens) were used to compare species. Our excavations reveal that burrow systems of Gunnison's prairie dog vary from simple to complex structures, with >50% of burrow systems having multiple aboveground entrances. There was no significant difference among species in total length of burrow. Burrow systems of black-tailed prairie dogs were significantly deeper than burrow systems of Gunnison's prairie dog. On average, burrow systems of Utah and Gunnison's prairie dogs had more aboveground entrances than black-tailed or white-tailed prairie dogs. Overall, the burrow systems for all species were similar in structure, complexity, and underground...

Increased risk of infectious disease is assumed to be a major cost of group living, yet empirical evidence for this effect is mixed. We studied whether larger social groups are more subdivided structurally. If so, the social subdivisions that form in larger groups may act as barriers to the spread of infection, weakening the association between group size and infectious disease. To investigate this ‘social bottleneck’ hypothesis, we examined the association between group size and four network structure metrics in 43 vertebrate and invertebrate species. We focused on metrics involving modularity, clustering, distance and centralization. In a meta-analysis of intraspecific variation in social networks, modularity showed positive associations with network size, with a weaker but still positive effect in cross-species analyses. Network distance also showed a positive association with group size when using intraspecific variation. We then used a theoretical model to explore the effects of subgrouping relative to other effects that influence disease spread in socially structured populations. Outbreaks reached higher prevalence when groups were larger, but subgrouping reduced prevalence. Subgrouping also acted as a ‘brake’ on disease spread between groups. We suggest research directions to understand the conditions under which larger groups become more subdivided, and to devise new metrics that account for subgrouping when investigating the links between sociality and infectious disease risk.

Although there is ample evidence for the generality of foraging and predation trade-offs in aquatic systems, its application to terrestrial systems is less comprehensive. In this review, meta-analysis was used to analyze experiments on giving-up-densities in terrestrial systems to evaluate the overall magnitude of predation risk on foraging behavior and experimental conditions mediating its effect. Results indicate a large and significant decrease in foraging effort as a consequence of increased predation risk. Whether experiments were conducted under natural or artificial conditions produced no change in the overall effect predation had on foraging. Odor and live predators as a correlate of predation risk had weaker and nonsignificant effects compared to habitat characteristics. The meta-analysis suggests that the effect of predation risk on foraging behavior in terrestrial systems is strongly dependent on the type of predation risk being utilized.

Group living in animals is believed to confer advantages related to a decrease in predation risk and an energetic trade-off between vigilance and foraging efficiency. Eight Gunnison's prairie dog, Cynomys gunnisoni, colonies in Flagstaff, Arizona (elevation 2300 m), were studied from April to August 2000 to examine the adaptive significance of colonial living in the context of predation risk and antipredator behavioral strategies. Each colony was sampled once every 10 days for a period of 3 h. Upright and quadrepedal vigilance was recorded using scan samples. All predation events were recorded. Results suggest that vigilant behavior in Gunnison's prairie dogs is an antipredator strategy because the animals oriented more frequently towards the periphery of the colony while vigilant. Gunnison's prairie dogs engaged in posting, an upright bipedal posture, more frequently than scanning, a quadrepedal posture. Furthermore, there was no relationship between either form of vigilance and population size. The proportion of animals vigilant decreased significantly only on the two smallest colonies as colony size increased. On larger colonies there was no relationship between the proportion of animals vigilant and colony size. The lack of change in the proportion of animals vigilant in larger populations may be a function of perceived risk rather than actual individual risk.