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Summary1. Network analysis is widely used in diverse fields and can be a powerful framework for studying the structure of biological systems. Temporal dynamics are a key issue for many ecological and evolutionary questions. These dynamics include both changes in network topology and flow on the network. Network analyses that ignore or do not adequately account for the temporal dynamics can result in inappropriate inferences. 2. We suggest that existing methods are currently under-utilized in many ecological and evolutionary network analyses and that the broader incorporation of these methods will considerably advance the current field. Our goal is to introduce ecologists and evolutionary biologists interested in studying network dynamics to extant ideas and methodological approaches, at a level appropriate for those new to the field. 3. We present an overview of time-ordered networks, which provide a framework for analysing network dynamics that addresses multiple inferential issues and permits novel types of temporally informed network analyses. We review available methods and software, discuss the utility and considerations of different approaches, provide a worked example analysis and highlight new research opportunities in ecology and evolutionary biology.

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Here, we present estimates of heritability and selection on network traits in a single population, allowing us to address the evolutionary potential of social behavior and the poorly understood link between sociality and fitness. To evolve, sociality must have some heritable basis, yet the heritability of social relationships is largely unknown. Recent advances in both social network analyses and quantitative genetics allow us to quantify attributes of social relationships and estimate their heritability in free-living populations. Our analyses addressed a variety of measures (in-degree, out-degree, attractiveness, expansiveness, embeddedness, and betweenness), and we hypothesized that traits reflecting relationships controlled by an individual (i.e., those that the individual initiated or were directly involved in) would be more heritable than those based largely on the behavior of conspecifics. Identifying patterns of heritability and selection among related traits may provide insight into which types of relationships are important in animal societies. As expected, we found that variation in indirect measures was largely explained by nongenetic variation. Yet, surprisingly, traits capturing initiated interactions do not possess significant additive genetic variation, whereas measures of received interactions are heritable. Measures describing initiated aggression and position in an agonistic network are under selection (0.3 < |S| < 0.4), although advantageous trait values are not inherited by offspring. It appears that agonistic relationships positively influence fitness and seemingly costly or harmful ties may, in fact, be beneficial. Our study highlights the importance of studying agonistic as well as affiliative relationships to understand fully the connections between sociality and fitness.animal model | animal social networks | yellow-bellied marmots B ehavioral ecologists have long viewed sociality and social relationships as adaptive traits shaped by evolution (1, 2). However, if we are to study the evolution of sociality and social relationships, there must be heritable variation in traits describing individual social behavior. Numerous studies have identified heritable variation in animal dispositions (3, 4), morphological characteristics, and behavioral traits (5) that may affect how individuals interact with conspecifics, yet the role of genetics in social interactions themselves is poorly understood. If traits affecting social interactions are heritable, we may expect measures of social relationships to be explained somewhat by additive genetic factors.There has been a recent upsurge in using animal social networks as tools for studying the ecology, evolution, and adaptive significance of sociality (6-8). Networks are based on interactions between individuals, and a variety of measures have been developed to quantify how connected individuals are with others in the group (9). Although studies of nonhuman species have explored the development of social networks (10) as well as the causes (11-13) and...

Individuals frequently leave home before reaching reproductive age, but the proximate causes of natal dispersal remain relatively unknown. The social cohesion hypothesis predicts that individuals who engage in more (affiliative) interactions are less likely to disperse. Despite the intuitive nature of this hypothesis, support is both limited and equivocal. We used formal social network analyses to quantify precisely both direct and indirect measures of social cohesion in yellow-bellied marmots. Because approximately 50 per cent of female yearlings disperse, we expected that social relationships and network measures of cohesion would predict dispersal. By contrast, because most male yearlings disperse, we expected that social relationships and cohesion would play a less important role. We found that female yearlings that interacted with more individuals, and those that were more socially embedded in their groups, were less likely to disperse. For males, social interactions were relatively unimportant determinants of dispersal. This is the first strong support for the social cohesion hypothesis and suggests that the specific nature of social relationships, not simply the number of affiliative relationships, may influence the propensity to disperse.

Social structure is a fundamental component of a population that drives ecological and evolutionary processes ranging from parasite transmission to sexual selection. Nevertheless, we have much to learn about factors that explain variation in social structure. We used advances in biologging and social network analysis to experimentally test how the local habitat, and specifically habitat complexity, modulates social structure at different levels in wild populations. Sleepy lizards, Tiliqua rugosa, establish nonrandom social networks that are characterized by avoidance of some neighbours and frequent interactions with one opposite-sex individual. Using synchronous GPS locations of all adult lizards, we constructed social networks based on spatial proximity of individuals. We increased habitat structural complexity in two study populations by adding 100 short fences across the landscape. We then compared the resulting movement behaviour and social structure between these populations and two unmanipulated populations. Social connectivity (network density) and social stability, measured at weekly intervals, were greater in populations with increased habitat structural complexity. The level of agonistic interaction (quantified as scale damage) was also higher, indicating a fitness cost of greater social connectivity. However, some network parameters were unaffected by increased complexity, including disassortative mixing by sex, and at the individual level, social differentiation among associates (coefficient of variation of edge weights) and maximal interaction frequencies (maximal edge weight). This suggests divergent effects of changed ecological conditions on individual association behaviour compared to the resulting social structure of the population. Our results contrast with those from studies of more gregarious species, in which higher structural complexity in the environment relaxed the social connectivity. This shows that the response to altered ecological conditions can differ fundamentally between species or between populations, and we suggest that it depends on their tendency for gregarious behaviour.

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