Potential banana skins in animal social network analysis

Social network analysis is an increasingly popular tool for the study of the fine-scale and global social structure of animals. It has attracted particular attention by those attempting to unravel social structure in fission–fusion populations. It is clear that the social network approach offers some exciting opportunities for gaining new insights into social systems. However, some of the practices which are currently being used in the animal social networks literature are at worst questionable and at best over-enthusiastic. We highlight some of the areas of method, analysis and interpretation in which greater care may be needed in order to ensure that the biology we extract from our networks is robust. In particular, we suggest that more attention should be given to whether relational data are representative, the potential effect of observational errors and the choice and use of statistical tests. The importance of replication and manipulation must not be forgotten, and the interpretation of results requires care. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Structure of the social network and its influence on transmission dynamics in a honeybee colony

Infectious processes in a social group are driven by a network of contacts that is generally structured by the organization arising from behavioral and spatial heterogeneities within the group. Although theoretical models of transmission dynamics have placed an overwhelming emphasis on the importance of understanding the network structure in a social group, empirical data regarding such contact structures are rare. In this paper, I analyze the network structure and the correlated transmission dynamics within a honeybee colony as determined by food transfer interactions and the changes produced in it by an experimental manipulation. The study demonstrates that widespread transmission in the colony is correlated to a lower clustering coefficient and higher robustness of the social network. I also show that the social network in the colony is determined by the spatial distribution of various age classes, and the resulting organizational structure provides some amount of immunity to the young individuals. The results of this study demonstrates how, using the honeybee colony as a model system, concepts in network theory can be combined with those in behavioral ecology to gain a better understanding of social transmission processes, especially those related to disease dynamics.     

Structure and resilience of the social network in an insect colony as a function of colony size

Social interactions are critical to the organization of worker activities in insect colonies and their consequent ecological success. The structure of this interaction network is therefore crucial to our understanding of colony organization and functioning. In this paper, I study the properties of the interaction network in the colonies of the social wasp Ropalidia marginata. I find that the network is characterized by a uniform connectivity among individuals with increasing heterogeneity as colonies become larger. Important network parameters are found to be correlated with colony size and I investigate how this is reflected in the organization of work in colonies of different sizes. Finally, I test the resilience of these interaction networks by experimental removal of individuals from the colony and discuss the structural properties of the network that are related to resilience in a social network. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau, and R. James).     

Cyclicity in the structure of female baboon social networks

There is an established and very influential view that primate societies have identifiable, persistent social organizations. It assumes that association patterns reflect long-term strategic interests that are not qualitatively perturbed by short-term environmental variability. We used data from two baboon troops in markedly different habitats over three consecutive seasons to test this assumption. Our results demonstrate pronounced cyclicity in the extent to which females maintained differentiated relationships. When food was plentiful, the companionships identified by social network analysis in the food-scarce season disappeared and were replaced by casual acquaintanceships more representative of mere gregariousness. Data from the fourth, food-scarce, season at one site indicated that few companions were re-united. It is likely that this reflected stochastic variation in individual circumstances. These results suggest that attention could profitably be paid to the effects of short-term local contingencies on social dynamics, and has implications for current theories of primate cognitive evolution. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Social network theory in the behavioural sciences: potential applications

Social network theory has made major contributions to our understanding of human social organisation but has found relatively little application in the field of animal behaviour. In this review, we identify several broad research areas where the networks approach could greatly enhance our understanding of social patterns and processes in animals. The network theory provides a quantitative framework that can be used to characterise social structure both at the level of the individual and the population. These novel quantitative variables may provide a new tool in addressing key questions in behavioural ecology particularly in relation to the evolution of social organisation and the impact of social structure on evolutionary processes. For example, network measures could be used to compare social networks of different species or populations making full use of the comparative approach. However, the networks approach can in principle go beyond identifying structural patterns and also can help with the understanding of processes within animal populations such as disease transmission and information transfer. Finally, understanding the pattern of interactions in the network (i.e. who is connected to whom) can also shed some light on the evolution of behavioural strategies.     

Social networks and non-market valuations

This paper considers the role of social networks in the non-market valuation of public goods. In the model individuals derive utility both from their own direct enjoyment of the public good and from the enjoyment of those in their network. We find that network structure almost always matters, both for utility and for valuation. The network increases aggregate valuation when it assigns higher importance, that is, stronger connections, to individuals with higher private values for the public good. The model provides a theoretical foundation for the idea of opinion leaders who have disproportionate influence over their communities. Specifically, opinion leaders are individuals assigned high importance by the network, and projects favored by opinion leaders tend to be favored by the network as a whole. The model can also guide future empirical studies by enabling a more structural approach to non-market valuation in a socially connected group.     

Food exploitation by a winter flock of greylag geese: behavioral dynamics,competition and social status

Summary To investigate the dynamics of the winter flock patch exploitation, feeding experiments were performed with 140 semi-tame, free-roaming greylag geese (Anser anser). Three different initial densities of barley were offered on a 50-m² patch: low (1600 grains/m²), intermediate (3300/m²) and high (I 1500/m²). Goose numbers on the patch, peck rates, and frequencies of agonistic encounters and of alert postures were observed by scanning the flock and in focal individuals. At low and intermediate initial food densities, peck rates decreased with food density, whereas at high food density, peck rates decreased only slightly over the feeding bout. Agonistic interactions increased as food decreased. A switch from exploitation to interference competition occurred at a threshold of approximately 900 remaining grains per square meter. With high initial food density, agonistic encounters began to increase gradually after 20–30 min. The same general patterns were observed for frequency of alertness. Family members occupied the patch for the longest time periods. The family female and the offspring fed most intensely. The family gander however, fed significantly less but was more aggressive and vigilant than all other social categories. Solitary geese spent their time on the patch doing little else but feeding and were the first to leave. It seems that high-ranking families and low-ranking singles adopt different competitive strategies, the former being superior interference competitors whereas the latter are forced scramble competitors.This study is dedicated to the memory of Konrad Lorenz Correspondence to: K. Kotrschal     

The social network structure of a wild meerkat population: 1. Inter-group interactions

Groups of individuals frequently interact with each other, but typically analysis of such interactions is restricted to isolated dyads. Social network analysis (SNA) provides a method of analysing polyadic interactions and is used to analyse interactions between individuals. We use a population of 12 groups (ca. 250 animals) of wild meerkats (Suricata suricatta) to test whether SNA can also be used to describe and elucidate patterns of inter-group interactions. Using data collected over 24 months, we constructed two sets of networks, based on direct encounters between groups and instances of roving males visiting other groups. We analysed replicated networks of each type of interaction to investigate similarities between networks of different social interactions as well as testing their stability over time. The two network types were similar to each other when derived from long-term data, but showed significant differences in structure over shorter timescales where they varied according to seasonal and ecological conditions. Networks for both types of inter-group interaction constructed from data collected over 3 months reliably described long-term (12- and 24-month) patterns of interactions between groups, indicating a stable social structure despite variation in group sizes and sex ratios over time. The centrality of each meerkat group in roving interactions networks was unaffected by the sex ratio of its members, indicating that male meerkats preferentially visit geographically close groups rather than those containing most females. Indeed, the strongest predictors of network structure were spatial factors, suggesting that, in contrast to analyses of intra-group interactions, analyses of inter-group interactions using SNA must take spatial factors into account.     

SOCPROG programs: analysing animal social structures

SOCPROG is a set of programs which analyses data on animal associations. Data usually come from observations of the social behaviour of individually identifiable animals. Associations among animals, sampling periods, restrictions on the data and association indices can be defined very flexibly. SOCPROG can analyse data sets including 1,000 or more individuals. Association matrices are displayed using sociograms, principal coordinates analysis, multidimensional scaling and cluster analyses. Permutation tests, Mantel and related tests and matrix correlation methods examine hypotheses about preferred associations among individuals and classes of individual. Weighted network statistics are calculated and can be tested against null hypotheses. Temporal analyses include displays of lagged association rates (rates of reassociation following an association). Models can be fitted to lagged association rates. Multiple association measures, including measures produced by other programs such as genetic or range use data, may be analysed using Mantel tests and principal components analysis. SOCPROG also performs mark-recapture population analyses and movement analyses. SOCPROG is written in the programming language MATLAB and may be downloaded free from the World Wide Web.     

Divided destinies: group choice by female savannah baboons during social group fission

Group living provides benefits to individuals while imposing costs on them. In species that live in permanent social groups, group division provides the only opportunity for nondispersing individuals to change their group membership and improve their benefit to cost ratio. We examined group choice by 81 adult female savannah baboons (Papio cynocephalus) during four fission events. We measured how each female’s group choice was affected by several factors: the presence of her maternal kin, paternal kin, age peers, and close social partners, her average kinship to groupmates, and her potential for improved dominance rank. Maternal kin, paternal kin, and close social partners influenced group choice by some females, but the relative importance of these factors varied across fissions. Age peers other than paternal kin had no effect on group choice, and average kinship to all groupmates had the same effect on group choice as did maternal kin alone. Most females were subordinate to fewer females after fissions than before, but status improvement did not drive female group choice; females often preferred to remain with social superiors who were their close maternal kin, rather than improving their own social ranks. We suggest that during permanent group fissions, female baboons prefer to remain with close maternal kin if those are abundant enough to influence their fitness; if they have too few close maternal kin then females prefer to remain with close paternal kin, and social bonds with nonkin might also become influential. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Association networks in spider monkeys (<Emphasis Type="Italic">Ateles geoffroyi</Emphasis>)

We use two novel techniques to analyze association patterns in a group of wild spider monkeys (Ateles geoffroyi) studied continuously for 8 years. Permutation tests identified association rates higher or lower than chance expectation, indicating active processes of companionship and avoidance as opposed to passive aggregation. Network graphs represented individual adults as nodes and their association rates as weighted edges. Strength and eigenvector centrality (a measure of how strongly linked an individual is to other strongly linked individuals) were used to quantify the particular role of individuals in determining the network's structure. Female–female dyads showed higher association rates than any other type of dyad, but permutation tests revealed that these associations cannot be distinguished from random aggregation. Females formed tightly linked clusters that were stable over time, with the exception of immigrant females who showed little association with any adult in the group. Eigenvector centrality was higher for females than for males. Adult males were associated mostly among them, and although their strength of association with others was lower than that of females, their association rates revealed a process of active companionship. Female–male bonds were weaker than those between same-sex pairs, with the exception of those involving young male adults, who by virtue of their strong connections both with female and male adults, appear as temporary brokers between the female and male clusters of the network. This analytical framework can serve to develop a more complete explanation of social structure in species with high levels of fission–fusion dynamics. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James)     

Females go where the food is: does the socio-ecological model explain variation in social organisation of solitary foragers?

The socio-ecological model (SEM) links ecological factors with characteristics of social systems and allows predictions about the relationships between resource distribution, type of competition and social organisation. It has been mainly applied to group-living species but ought to explain variation in social organisation of solitary species as well. The aim of this study was to test basic predictions of the SEM in two solitary primates, which differ in two characteristics of female association patterns: (1) spatial ranging and (2) sleeping associations. Beginning in August 2002, we regularly (re-)captured and marked individuals of sympatric populations of Madame Berthe's and grey mouse lemurs (Microcebus berthae, Microcebus murinus) in Kirindy Forest (Madagascar). We recorded data on spatial patterns, feeding and social behaviour by means of direct observation of radio-collared females. The major food sources of M. berthae occurred in small dispersed patches leading to strong within-group scramble competition and over-dispersed females with a low potential for female associations. In contrast, M. murinus additionally used patchily distributed, high-quality (large) resources facilitating within-group contest competition. The combined influence of less strong within-group scramble and contest as well as between-group contest over non-food resources allowed females of this species to cluster in space. Additionally, we experimentally manipulated the spatial distribution of food sources and found that females adjusted their spatial patterns to food resource distribution. Thus, our results support basic predictions of the SEM and demonstrated that it can also explain variation in social organisation of solitary foragers.     

Small queens in the ant Ectatomma tuberculatum: a new case of social parasitism

Social parasites exploit the worker force of colonies of other social insects to rear their own young. Social parasitism occurs in several Hymenoptera and is particularly common in several tribes of the ant subfamilies Myrmicinae and Formicinae. Here, we document the occurrence of miniaturized queens (microgynes) in colonies of Ectatomma tuberculatum, an ant belonging to the subfamily Ectatomminae. Behavioral observations and genetic analyses show that microgynes concentrate their reproductive efforts almost exclusively on the production of sexual offspring (microgynes and males), whereas the regular, large queens (macrogynes) produce workers in addition to sexuals. According to mitochondrial and nuclear markers, gene flow between microgynes and macrogynes is extremely limited. Whereas the co-occurrence of microgynes and macrogynes in the related species Ectatomma ruidum constitutes an intraspecific polymorphism associated with alternative dispersal tactics, microgynes found in colonies of E. tuberculatum appear to be a distinct species and to represent the first case of social parasitism in the poneromorph subfamilies of ants.     

Genetic population structure and sociogenetic organisation in Formica truncorum (Hymenoptera; Formicidae)

Summary The genetic population structure and the sociogenetic organization of the red wood ant Formica truncorum were compared in two populations with monogynous colonies and two populations with polygynous colonies. The genetic population structure was analysed by measuring allele frequency differences among local subsets of the main study populations. The analysis of sociogenetic organisation included estimates of nestmate queen and nestmate worker relatedness, effective number of queens, effective number of matings per queen, relatedness among male mates of nestmate queens and relatedness between queens and their male mates. The monogynous populations showed no differentiation between subpopulations, whereas there were significant allele frequency differences among the subpopulations in the polygynous population. Workers, queens and males showed the same genetical population structure. The relatedness among nestmate workers and among nestmate queens was identical in the polygynous societies. In three of the four populations there was a significant heterozygote excess among queens. The queens were related to their male mates in the polygynous population analysed, but not in the monogynous ones. The data suggest limited dispersal and partial intranidal mating in the populations with polygynous colonies and outbreeding in the populations having monogynous colonies. Polyandry was common in both population types; about 50% of the females had mated at least twice. The males contributed unequally to the progeny, one male fathering on average 75% of the offspring with double mating and 45–80% with three or more matings. Correspondence to: L. Sundström     

论近年来畜禽疫病的特点及其预防对策

随着畜牧业的快速发展,大规模、高密度、集约化的饲养和频繁的调运等,使畜禽更易发生流行性、群发性的疫病.畜禽疫病造成的损失影响了人类的生活需要,而且某些人畜共患的疫病还给人类健康带来了严重的威胁.要保证畜牧业可持续发展和保护人类健康,控制畜禽疫病就是关健措施.由于近年畜禽疫病的变化,我们的防治对策应发生相应变化.     

Social structure and co-operative interactions in a wild population of guppies (Poecilia reticulata)

In contrast to the substantial number of theoretical papers that have examined the mechanisms by which cooperation may evolve, very few studies have investigated patterns of co-operation in natural animal populations. In the current study, we use a novel approach, social network analysis, to investigate the structure of co-operative interactions in the context of predator inspection in a wild population of guppies (Poecilia reticulata). Female guppies showed social preferences for stable partners, fulfilling a key assumption made by models of reciprocity. In the laboratory, wild female guppies disproportionately engaged in predator inspection with others with whom they had strong social associations. Furthermore, pairs of fish that frequently engaged in predator inspection did so in a particularly co-operative way, potentially reducing costs associated with predator inspection. Taken together, these results provide evidence for assortative interactions forming the basis of co-operation during predator inspection in a natural fish population. The occurrence of highly interconnected social networks between stable partners suggests the existence of co-operation networks in free-ranging populations of the guppy.     

Plasticity of social organization in a forest ant species

We manipulated availability of food and nesting sites in one population of the forest ant Myrmica punctiventris. The manipulations produced significant changes in relatedness structure, reproductive allocation, and response to hierarchical selection. Food availability appeared to have a consistently stronger influence on these aspects of social organization than did availability of nesting sites. We interpret our experimental results in light of observed differences between populations, and discuss implications for kin selection dynamics. Received: 30 July 1998 / Accepted after revision: 31 October 1998     

Structural path analysis of ecosystem networks

This paper explains the application of structural path analysis (SPA), an input–output-based technique for measuring flows in ecological and linked ecological–economic networks. Previous methods of input–output flow analysis have concluded with aggregate indexes relying on the summing feature of the Leontief inverse in order to completely account for throughflows along a multitude of inter-compartmental paths. This paper shows that for most linear dissipative networks, a manageable number of paths of limited length exist that cover in the order of 99% or more of total throughflow. These paths can be conveniently extracted, enumerated and ranked using SPA.     

Combining state and transition models with dynamic Bayesian networks

Bashari et al. (2009) propose combining state and transition models (STMs) with Bayesian networks for decision support tools where the focus is on modelling the system dynamics. There is already an extension of Bayesian networks - so-called dynamic Bayesian networks (DBNs) - for explicitly modelling systems that change over time, that has also been applied in ecological modelling. In this paper we propose a combination of STMs and DBNs that overcome some of the limitations of Bashari et al.’s approach including providing an explicit representation of the next state, while retaining its advantages, such an the explicit representation of transitions. We then show that the new model can be applied iteratively to predict into the future consistently with different time frames. We use Bashari et al.’s rangeland management problem as an illustrative case study. We present a comparative complexity analysis of the different approaches, based on the structure inherent in the problem being modelled. This analysis showed that any models that explicitly represent all the transitions only remain tractable when there are natural constraints in the domain. Thus we recommend modellers should analyse these aspects of their problem before deciding whether to use the framework.     

The influence of social relationships on pro-environment behaviors

We examine how social relationships are related to pro-environment behaviors. We use new data from a nationally representative US sample to estimate latent cluster models in which we describe individuals' profiles of social ties with family, neighbor, and coworkers along two dimensions: intensity of connections and pro-environment norms. While our results confirm the link between social ties and economic behaviors, we show that ties among relatives, neighbors, and coworkers are not perfect substitutes. In particular, we observe consistent relationships between green family profiles and altruistic and community-based behaviors. We also find that the effect of coworker ties is visible for cost-saving activities and altruistic behaviors, and that neighbors matter for working with others in the community to solve a local problem, volunteering, and recycling.