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Hasenjager, M. J., & Dugatkin, L. A. Social Network Analysis in Behavioral Ecology. Advances in the Study of Behavior. Academic Press.
Abstract: Abstract In recent years, behavioral ecologists have embraced social network analysis (SNA) in order to explore the structure of animal societies and the functional consequences of that structure. We provide a conceptual introduction to the field that focuses on historical developments, as well as on novel insights generated by recent work. First, we discuss major advances in the analysis of nonhuman societies, culminating in the use of SNA by behavioral ecologists. Next, we discuss how network-based approaches have enhanced our understanding of social structure and behavior over the past decade, focusing on: (1) information transmission, (2) collective behaviors, (3) animal personality, and (4) cooperation. These behaviors and phenomena possess several features—e.g., indirect effects, emergent properties—that network analysis is well equipped to handle. Finally, we highlight recent developments in SNA that are allowing behavioral ecologists to address increasingly sophisticated questions regarding the structure and function of animal sociality.
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Baragli, P., Cozzi, A., Rugani, R., Sighieria, C., & Regolin, L. (2008). Delayed search for non-social goals by Equids (Equus caballus and Equus asinus). In IESM 2008.
Abstract: Delayed-responses have been traditionally employed to investigate the temporal characteristics of animals“ ability to represent and recall objects that have disappeared. In the typical condition, the animal, usually a mammal, observes the experimenter hiding an interesting goal (e.g. some food) in a certain location. A delayed-response task (DRT) was administered to 4 female Esperia pony (2 years old) coming from a free-range breed (Frosinone, Italy) and to 7 female Amiata donkeys (4.2±2 years old) coming from a conservation stock (University of Pisa, Italy). The DRT's apparatus was located in a square fence. A single ”U-shaped“ screen (330x160x140 cm) made by wood shavings blocks was positioned in the centre of the fence. A gap (40x50 cm) on the ground was in the middle of the central side of the U-shaped-screen and served to make the food-attractor disappear. The food-attractor consisted in cereal flakes and fresh grass for ponies and cereal flakes for donkeys. A bucket full of food was placed on a dolly tied on a rope which could be pulled by an experimenter. In a preliminary training each animal was allowed to eat food from the bucket and, while the animal was eating, the dolly was gently pulled away from the animal, and beyond the screen through the gap. The subjects needed to move around of the screen in order to retrieve the food. As a reinforcement, they were allowed to eat some food from the bucket once behind the screen. From trial to trial, the bucket was presented farther and farther (starting with a distance of 1 m in front of the screen to reach 7 m). Therefore subjects were tested in the DRT requiring them to rejoin the bucket with the goal-food disappearing behind the screen as in the preliminary training but following a 10 s delay. For the DRT, the bucket was placed 7 m in front of the screen, 3 m away from the animal's starting area. Then the dolly was pulled away from the animal. Ten seconds after the disappearance of the dolly behind the screen the animal was released from the starting area. The DRT ended when the subject had reached the attractor behind the screen on 3 consecutive trials. Results showed that all animals were able to rejoin the food behind the screen after 10 s delay. The mean time of the delayed-response (mean±sd, in s) in the ponies (1st: 19.8±8; 2nd: 10.8±2.2; 3rd: 12.8±2.8) and in the donkeys (1st: 28.4±10; 2nd: 26.9±13; 3rd: 24.3±16.6) showed a trend to decrease from first trial to third. These preliminary results suggest that like other mammals our ponies and donkeys can maintain a working memory trace of the location where biologically attractive objects have been seen to disappear. In conclusion, this study paves the way to set up a viable model system for the investigation of the more sophisticated aspects of Equids” cognitive abilities such as working memory.
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P. K. McGregor,, & T. M. Peake,. (2000). Communication networks: social environments for receiving and signalling behaviour. Acta. Ethol., 2(2), 71–81.
Abstract: Communication and social behaviour are inextricably linked, with communication mediating important social behaviours such as resource defence and mate attraction. However, the social environment in which communication occurs is often ignored in discussions of communication behaviour. We argue that networks of several individuals are the common social environment for communication behaviour. The consequences for receivers and signallers of communicating in a network environment are the main subjects of this review. Eavesdropping is a receiving behaviour that is only possible in the environment of a network and therefore we concentrate on this behaviour. The main effect of communication networks on signallers is to create competition with other signallers for receiver attention. We discuss the consequences of such competition. To conclude, we explore the role of signals and signalling interactions as sources of information that animals exploit to direct their behaviour.
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Crystal, J. D. (1999). Systematic nonlinearities in the perception of temporal intervals. J Exp Psychol Anim Behav Process, 25(1), 3–17.
Abstract: Rats judged time intervals in a choice procedure in which accuracy was maintained at approximately 75% correct. Sensitivity to time (d') was approximately constant for short durations 2.0-32.0 s with 1.0- or 2.0-s spacing between intervals (n = 5 in each group, Experiment 1), 2.0-50.0 s with 2.0-s spacing (n = 2, Experiment 1), and 0.1-2.0 s with 0.1- or 0.2-s spacing (n = 6 in each group, Experiment 2). However, systematic departures from average sensitivity were observed, with local maxima in sensitivity at approximately 0.3, 1.2, 10.0, 24.0, and 36.0 s. Such systematic departures from an approximately constant d' are predicted by a connectionist theory of time with multiple oscillators and may require a modification of the linear timing hypothesis of scalar timing theory.
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Wey, T., Blumstein, D. T., Shen, W., & Jordán, F. (2008). Social network analysis of animal behaviour: a promising tool for the study of sociality. Anim. Behav., 75(2), 333–344.
Abstract: Social animals live and interact together, forming complex relationships and social structure. These relationships can have important fitness consequences, but most studies do not explicitly measure those relationships. An approach that explicitly measures relationships will further our understanding of social complexity and the consequences of both direct and indirect interactions. Social network analysis is the study of social groups as networks of nodes connected by social ties. This approach examines individuals and groups in the context of relationships between group members. Application of social network analysis to animal behaviour can advance the field by identifying and quantifying specific attributes of social relationships, many of which are not captured by more common measures of sociality, such as group size. Sophisticated methods for network construction and analysis exist in other fields, but until recently, have seen relatively little application to animal systems. We present a brief history of social network analysis, a description of basic concepts and previous applications to animal behaviour. We then highlight relevance and constraints of some network measures, including results from an original study of the effect of sampling on network parameter estimates, and we end with promising directions for research. By doing so, we provide a prospective overview of social network analysis' general utility for the study of animal social behaviour.
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Whishaw, I. Q., Sacrey, L. - A. R., & Gorny, B. (2009). Hind limb stepping over obstacles in the horse guided by place-object memory. Behav. Brain. Res., 198(2), 372–379.
Abstract: An animal that has stepped over an obstacle with its forelimbs uses a memory of the obstacle to guide the hind limbs so that they also clear the obstacle, even in situations in which long pauses are introduced between forelimb and hind limb stepping. To further clarify the features of hind limb obstacle clearance memory, the present study examined hind limb obstacle clearance in the horse. A rider guided horses over obstacles and paused the horse over obstacles in tests that examined the relationship between forelimb and hind limb stepping, with the following results. First, the horses displayed memory for an obstacle as measured by hind limb lifting over the obstacle for durations lasting as long as 15Â min. The response was not dependent upon ongoing visualization of the obstacle, as limb lifting was unaffected by visual occlusion with blinders, a blindfold, or by removing the obstacle during the pause. Second, previous experience of stepping over an obstacle led to pause-related hind limb lifting at the object's previous location even on trials for which there was no obstacle and so no preceding forelimb lifting. Third, whereas a horse would lift its hind limbs to clear two successively presented obstacles, replacing an obstacle before the horse after the forelimbs had cleared the obstacle prevented subsequent hind limb lifting at the obstacle's previous location. Taken together the results show that hind limb obstacle clearance is guided by a place-object memory. The results are discussed in relation to the differential sensory and memonic control of forelimb and hind limb stepping with the suggestion that place-object memory can guide hind stepping as well as overshadow working memory from front leg stepping.
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VanDierendonck, M. C., de Vries, H., Schilder, M. B. H., Colenbrander, B., & Þorhallsdóttir, A. G. and S., H. (2009). Interventions in social behaviour in a herd of mares and geldings. Appl. Anim. Behav. Sci., 116(1), 67–73.
Abstract: Social dynamics and maintenance of social cohesion were studied by analysing social interventions in two groups of horses consisting of adult mares, their offspring, adult geldings and sub-adults. The animals were observed for a total of 1316 h. All relevant dyadic and triadic social interactions, including initial behaviour, possible intervention and outcome, were recorded. The main question was: do horses use interventions in affiliative interactions to safeguard their social network? Horses were significantly more likely to intervene in allogrooming or play interactions that involved a preferred partner. The stronger the preferred association in allogrooming, the higher the likelihood the intervener took over allogrooming with an initial dyad member. Interveners originating from two newly introduced groups (n = 3 and 5), intervened significantly more often when a member of their own group allogroomed with an unfamiliar horse. In play, no correlation with unfamiliarity was found. Overall, the intervening horses stopped more than half of the initial allogrooming interactions, and a third of all interactions. Therefore, social facilitation cannot sufficiently explain interference behaviour. This study shows that maintaining relationships with preferred partners is important to horses and has implications for equine husbandry and management.
Keywords: Horses; Social relationships; Interventions; Mares; Geldings; Play; Allogrooming; Social network
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Sueur, C., & Petit, O. (2008). Organization of Group Members at Departure Is Driven by Social Structure in Macaca. Int. J. Primatol., 29(4), 1085–1098.
Abstract: Abstract Researchers have often explained order of progression of group members during joint movement in terms of the influence of ecological pressures but rarely that of social constraints. We studied the order of joining by group members to a movement in semifree-ranging macaques with contrasting social systems: 1 group of Tonkean macaques (Macaca tonkeana) and 1 group of rhesus macaques (M. mulatta). We used network metrics to understand roles and associations among individuals. The way the macaques joined a movement reflected the social differences between the species in terms of dominance and kinship. Old and dominant male rhesus macaques were more often at the front of the movement, contrary to the Tonkean macaques, which exhibited no specific order. Moreover, rhesus macaques preferred to join high-ranking or related individuals, whereas Tonkean macaques based associations during joining mostly on sexual relationships with a subgroup of peripheral males.
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Krause, J., Croft, D., & James, R. (2007). Social network theory in the behavioural sciences: potential applications. Behav. Ecol. Sociobiol., 62(1), 15–27.
Abstract: Abstract 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.
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Lusseau, D., Whitehead, H., & Gero, S. (2008). Incorporating uncertainty into the study of animal social networks. Anim. Behav., 75(5), 1809–1815.
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