Social Environment and Agonistic Interactions: Strategies in a Small Social Mammal

In group‐living species, the development of agonistic interactions among conspecifics may be affected by socio‐ecological factors, such as size and composition of social group, and availability of nests and food. We analysed the importance of size and composition of social groups on agonistic interactions among males in the Southern mountain cavy (Microcavia australis). We made behavioural observations in four social groups of different size and composition. We recorded two types of agonistic interactions: agonistic displays and direct agonistic behaviours; both types increased in the breeding season. A social group composed of a high number of males was associated with high frequency of agonistic displays. Direct agonistic behaviours were also influenced by the interaction of season and number of males per social group and number of females per social group. Agonistic interactions were also recorded among males of different socials groups in the breeding season. Agonistic displays were most frequent among males of the same social group, whereas direct agonistic behaviours were most common among males of different social groups. These results suggest that social factors affect agonistic interactions among males of Southern mountain cavy and that in a conflict situation, males develop different strategies, such as increased frequency of agonistic behaviours in breeding season and intragroup cooperation for defence of oestrous females.     

Effects of enrichment items on activity and social interactions in domestic horses (Equus caballus)

The aim of this study was to investigate the use of items intended to provide enrichment during turnout, both for individual and group kept horses in an attempt to reduce the amount of passive behaviours. The study was divided into two parts, where study 1 involved eight horses rotated through eight individual paddocks, each containing one of seven enrichment items and one paddock being kept without item, functioning as a control. The horses’ item-directed behaviours; passive behaviours or other non-item related activities were scored using instantaneous sampling, every minute for 1 h at the beginning and the end of the turnout period. Study 2 involved six horse groups (3–6 horses) and the same scoring methods and ethogram as in study 1. The four items that the horses interacted the most with during study 1 (straw STRA, ball filled with concentrates CBALL, branches BRAN and scratching pole POLE) are investigated in study 2. In addition, the amount of social interactions was recorded.Both horses kept individually (P < 0.05) and in groups (P < 0.0001) performed significantly more item-directed behaviours towards edible items like STRA and CBALL than other objects. There was, however, no overall relation between the numbers of item-directed behaviours and the number of passive behaviours observed, indicating that the enrichment items did not alone reduce the amount of passive behaviours during turnout periods. Such a reduction was, however, only apparent when horses spent more time eating green leaves growing on the paddock surface (R = ?0.97 study 1, R = ?0.67 study 2, P < 0.0001). Access to STRA in group kept horses also seemed to reduce the amount of agonistic behaviours (P < 0.0001). In conclusion, if grass is not available in paddocks, the provision of roughage reduces the amount of passive behaviours in singly kept horses and it also reduces the risk of agonistic interactions between horses kept in group.     

Grouping horses according to gender—Effects on aggression, spacing and injuries

Many horse owners tend to group horses according to gender, in an attempt to reduce aggressive interactions and the risk of injuries. The aim of our experiment was to test the effects of such gender separation on injuries, social interactions and individual distance in domestic horses. A total of 66 horses were recruited from 4 different farms in Norway and Denmark and divided into six batches. Within each batch, horses were allotted into one mare group, one gelding group and one mixed gender group, with most groups consisting of three or four animals. After 4–6 weeks of acclimatisation, a trained observer recorded all social interactions using direct, continuous observation 1 h in the morning and 1 h in the afternoon for three consecutive days. Recordings of the nearest neighbour of each horse were performed using instantaneous sampling every 10 min. The horses were inspected for injuries before grouping, day 1 after grouping and after 4–6 weeks. No significant effect of gender composition was found on social interactions (P > 0.05), spacing (P > 0.07) or injuries (P > 0.23). Eighty percent of all aggressive interactions recorded were threats, not involving physical contact. Horses with the smallest space allowance showed the highest mean number of aggressive interactions (28.6 ± 6.1 interactions per 6 h) compared to the mean of all the other batches (8.3 ± 1.0 interactions per 6 h). Very few injuries were found and most were superficial. In conclusion, gender composition does not seem to have any effect on aggression level, spacing or injuries. However, the early social experience of horses, management of feeding and space allowance probably represents more important factors for successful group housing of domestic horses.     

Keeping horses in groups: A review

Although husbandry conditions for horses have improved over the last decades, many horses are still kept singly with limited or no physical contact to other horses. This is surprising, given the fact that keeping horses in groups is recognised best to fulfil their physical and behavioural needs, especially their need for social contact with conspecifics, as well as to have a beneficial effect on horse–human interactions during training.Group housing of farm animals is widely applied in practice. As a consequence, scientists have investigated numerous aspects of group housing to help further improve animal welfare and human–animal interactions under these conditions. However, compared to this literature available in farm animals, and the plentiful studies conducted of feral horse populations, there is much less done when it comes to the management of horses kept in groups in the domestic environment. In particular, limited scientific information is available into the effect of group size and group composition on behaviour and methods of introducing new horses into established groups, even though problems related to social integration are repeatedly taken as arguments against keeping horses in groups.This review, therefore, aims to provide an overview of the current scientific knowledge regarding keeping horses in groups. Furthermore, it aims to give insight into whether or not some of the concerns related to keeping horses in groups are justified and to review scientifically based solutions that could be useful in practice to improve horse welfare and human safety.     

Identifying potential risk situations for humans when removing horses from groups

Removing a horse from its social group may be considered risky, both for the handler and the horse, because other horses can interfere in the catching process. The main aim of this study was to identify where and when these risk situations occur while removing a horse from its group. A potential risk situation was defined by the closeness of loose horses in the group or by any physical contact with them. Whether the number of horses following would be influenced by the social rank of the horse being led out, and whether more horses would follow to the gate when a larger proportion of the group was removed compared to when a single horse was taken out were also investigated.Thirty-two mares (1-2 years) were kept in groups of four. All horses were taken out of their home paddock twice alone (64 tests) and twice with a companion (32 tests). One handler (or two handlers when two horses were removed) was asked to approach (phase 1) and catch the target horse (phase 2), walk it to the centre of the paddock and remain stationary at a post for 30 s (phase 3), walk to the paddock entrance (phase 4) and through the gate (phase 5). The number of horses following, and the number of loose horses in proximity (<2 m, 2-5 m) to the target horse and handler was estimated, and horse-horse and horse-human interactions were recorded continuously for the five scoring phases.Significantly more loose horses were within 2 m of a single target horse during the phases approach (mean ± SD: 1.5 ± 0.8), catch (1.6 ± 0.9) and post (1.7 ± 0.7) than during walk (1.0 ± 0.5) and gate (1.1 ± 0.6). Rank did not influence the number of horses following to the gate (high rank: 2.4 ± 0.7; lower rank: 2.0 ± 1.0; P = 0.396) and interactions between horses were rare. A greater proportion of the loose horses followed when two horses (0.9 ± 0.2) were removed compared to when a single horse (0.7 ± 0.3) was taken out (P = 0.011).In conclusion, maintaining a distance to other horses in the group by reducing the time being relatively stationary, so giving loose horses fewer chances to approach, is likely to contribute to improved handler's safety. Removing a small proportion of the group may also decrease the probability of the other horses following.     

Relations between group size and composition and feeding competition in mountain gorilla groups

The intensity and patterning of interference competition for food within mountain gorilla (Pan gorilla beringei) social groups is influenced in several ways by group size and composition and varies within and across age/sex classes. Data on 251 spatial displacemtnts associated with feeding, collected during a 17-month study of mountain gorilla feeding ecology, show that overall displacement rates and displacement rates for individuals were positively correlated with social group size. Silverback males were responsible for a disproportionately high number of displacements. Adult females also were involved in competitive interactions over food more often than expected from their representation in groups, and had feeding bouts interrupted disproportionately often, principally by other females and by silverbacks. Competitive relationships between females varied in association with female dominance rank and age, but were not clearly associated with relatedness between females. The results support the argument that social foraging entails a cost which is proportional to group size and which falls particularly on adult females. The comparatively low rates of competitive interactions, however, suggest that this cost is relatively low, and that female mountain gorillas sacrifice little in terms of feeding efficiency by living in social groups.     

Attributes and validity of dominance hierarchy in the female pigtail macaque

A group of six unrelated female pigtail macaques,Macaca nemestrina, of the same age, was studied in captivity over a period of five years. The animals were observed under six different situations, following changes in the social composition of the group. The following social interactions were recorded: withdrawal, attack, threat, presentation, mount, and grooming. Although behavioural rates varied for each period, their distribution network was remarkably consistent. Each subject could be assigned a rank, which remained stable over the six periods. From an analysis of the number of dyads in which one of the two animal significantly performed both paired behaviours more than the other animal (external validity), behaviours clustered into two main groups: (1) attacks, threats, and withdrawals; and (2) mounts, presentations, and grooming. Quantitative methods were performed to standardize the degree in which linearity, stability, reciprocity, and idiosyncrasy of the interactions under study may account for social variability. When considering these properties, the behaviours clustered in the same two groups (“agonistic” and “affinitive”). A method to describe the dominance style ofMacaca nemestrina was proposed, which can easily be replicated for comparisons with females of other primate species and in different conditions.     

An equine protozoal myeloencephalitis challenge model testing a second transport after inoculation with Sarcocystis neurona sporocysts

Previous challenge studies performed at Ohio State University involved a transport-stress model where the study animals were dosed with Sarcocystis neurona sporocysts on the day of arrival. This study was to test a second transportation of horses after oral inoculation with S. neurona sporocysts. Horses were assigned randomly to groups: group 1, transported 4 days after inoculation (DAI); group 2, at 11 DAI; group 3, at 18 DAI; and group 4, horses were not transported a second time (controls). An overall neurologic score was determined on the basis of a standard numbering system used by veterinarians. All scores are out of 5, which is the most severely affected animal. The mean score for the group 1 horses was 2.42; group 2 horses was 2.5; group 3 horses was 2.75; and group 4 horses was 3.25. Because the group 4 horses did not have a second transport, they were compared with all other groups. Statistically different scores were present between group 4 and groups 1 and 2. There was no difference in the time of seroconversion between groups. There was a difference between the time of onset of first clinical signs between groups 1 and 4. This difference was likely because of the different examination days. Differences in housing and handling were likely the reason for the differences in severity of clinical signs. This model results in consistent, significant clinical signs in all horses at approximately the same time period after inoculation but was most severe in horses that did not experience a second transport.     

Direct and indirect effects of grazing constrain shrub encroachment in semi‐arid Patagonian steppes

Question: What are the long‐term effects of grazing exclusion on the population structure and dynamics of, and interactions among, three dominant shrub species? Location: Grass‐shrub Patagonian steppe, Chubut, Argentina. Methods: Permanent plots were established in grazed paddocks and paddocks excluded from grazing in representative Patagonian rangelands. Shrub abundance, population size‐structure, short‐term (two 3‐yr periods) and long‐term (matrix models) population dynamics, and neighborhood interactions of three native and codominant shrub species (Mulinum spinosum, Senecio filaginoides and Adesmia volckmanni) were measured and analysed using different statistical approaches. Results: The total density of shrubs was 74% higher in paddocks excluded from grazing, owing mainly to increases in Mulinum (80%) and Senecio (68%) species. However, differences in size structure between ungrazed and grazed paddocks were only detected in Mulinum. Demographic rates differed between shrub species, time‐periods and grazing conditions. In particular, recruitment in the short term (especially in wet years) and population growth rate in the long term (λ) were higher in paddocks excluded from grazing only in Mulinum populations. Senecio populations showed a marginal increase in recruitment and mortality independent of the grazing condition in the wet and dry period. Grazing exclusion modified the balance of neighborhood interactions among the three shrub species. In grazing‐exclusion paddocks, there was a balance between positive and negative interspecific interactions, while in grazed paddocks there were more negative intraspecific and interspecific interactions, resulting in a net negative balance of neighborhood interactions. Conclusions: Our understanding of woody encroachment in arid rangelands can be informed through evaluation of direct and indirect effects of grazing exclusion on the abundance and demography of dominant woody species. In Patagonian arid steppes, the occurrence of woody encroachment in rangelands excluded from grazing can be explained by altered responses in plant‐animal and plant‐plant interactions among shrub species.     

Contrasts in social structure among black-and-white colobus monkey groups

Three types of Colobus guereza groups may be distinguished on the bases of size and composition, namely small one-male groups, large, one-male groups and multi-male groups. The social structure of each type of group is described in terms of the distribution of non-agonistic interactions, the frequency and distribution of agonistic behaviour and the organization of the roles of vigilance, territorial defence and leadership. A number of differences are found between the group types which appear to be related to the differences in group size and composition. It is suggested that these group types represent stages in the life-cycle of colobus groups, and that such an interpretation may help to resolve some of the conflicting reports in the literature.     

Number of neighbors instead of group size significantly affects individual vigilance levels in large animal aggregations

The group size effect states that animals living in groups gain anti‐predator benefits through reducing vigilance levels as group size increases. A basic assumption of group size effect is that all individuals are equally important for a focal individual, who may adjust its vigilance levels according to social information acquired from them. However, some studies have indicated that neighbors pose greater influences on an individual's vigilance decisions than other group members, especially in large aggregations. Vigilance has also been found to be directed to both predators (anti‐predation vigilance) and conspecifics (social vigilance). Central individuals might rely more on social vigilance than peripheral individuals. To test these hypotheses, we examined the effects of flock size, number of neighbors and position within a flock on vigilance and competition of greater white‐fronted goose Anser albifrons that form large foraging flocks in winter, controlling the effects of other variables (group identity, winter period and site). We found that individual vigilance levels were significantly affected by number of neighbors and position within a flock, whereas flock size showed no effect. Individuals devoted a large component of vigilance to nearby flock mates. Central individuals directed a relatively larger proportion of vigilance to monitor neighbors than peripheral ones, indicating that central individuals more relied on social information acquired from neighbors, possibly caused by the more blocked visual field of central individuals. Moreover, some social vigilance may function as conducting or preventing agonistic interactions since competition intensity was positively correlated with number of neighbors. Our study therefore demonstrate that the number of neighbors is more important than group size in determining individual vigilance in large animal groups. Further studies are still needed to unravel which neighbors pose greater influence on individual vigilance, and the factors that influence individuals to acquire information from their neighbors to adjust vigilance behaviors.     

Preliminary investigation of social interactions and feeding behavior in captive group-housed Tasmanian devils (Sarcophilus Harrisii)

As the number of Tasmanian devils (Sarcophilus harrisii) in captivity increases, an understanding of captive social dynamics and behavior is becoming increasingly important. In the wild, devils are solitary, although sometimes, they congregate to feed on a large carcass. However, it is common to house devils in groups as a form of social enrichment. This study investigated how behavior at feeding time of captive Tasmanian devils varied in groups of different sizes. Observations were made of individually housed devils and devils in groups of two, three, five, and six, when presented with a carcass on which to feed. Total feeding duration ranged from 6.5 to 47.4 minutes per observation period (70 minutes). There was no significant interaction between feeding duration and group size during the experiment. Feeding duration varied daily and depended on carcass size. Social housing of Tasmanian devils enabled them to display dyadic and agonistic behaviors during feeding. Observing behaviors and learning from the outcomes of these interactions can improve husbandry techniques. Creating a captive environment that encourages natural behaviors may enhance survival in the wild following translocation.     

Contamination of surface run-off water and soil in two horse paddocks

Increased stabling of horses near to cities has led to interest in the environmental effects of paddocks. In this study, the contamination of horse paddocks was examined by determining the nutrient and micro-organism contents in the surface run-off waters and the electrical conductivity, pH and phosphorus, potassium and nitrate contents of top soils. Two open-stable paddocks were studied, one cleaned and the other left uncleaned, with a stocking density of 37.5 animalsha(-1) in both. The feeding and drinking places were the most contaminated areas of both paddocks. In spring, after seven months of use, the nutrient concentrations in the surface run-off water from paddocks were 3.4-18.8mg/l for total phosphorus, 3.0-15.0mg/l for phosphate and 18.3-140.0mg/l for total nitrogen, indicating a risk to surface waters. Summer rain generated surface run-off, especially from the feeding area of the stock-free uncleaned paddock.     

Effect of group size and caste ratio on individual survivorship and social immunity in a subterranean termite

Individuals living within social groups may benefit from the efficiencies of division of labour, but on the other hand render themselves vulnerable to socially transmitted disease. This cost to social living should promote cooperative barriers to disease transmission, especially in eusocial taxa where spatial and genetic proximity to nestmates are characteristically pronounced. Termites are eusocial yet little is known about how their sociality is deployed to resist contagion. In this study, we manipulate two variables that are expected to affect the number and nature of social interactions and measure the ability of individuals within groups to resist fungal infection. From laboratory experiments on field-collected colonies, we report that both group size and caste composition directly affect the survivorship of individuals within groups, but only caste composition moderates survivorship upon immune challenge. Our study therefore provides no statistical evidence that individual Eastern subterranean termites (Reticulitermes flavipes) have increased resistance to disease in crowded groups—that is, there is no evidence for a density-dependent social immune response. Our results do suggest, however, that the caste-specific nature of interactions may be important for controlling disease in a social context.     

Effects of social group size on information transfer and task allocation

Summary Social animals exchange information during social interaction. The rate of interaction and, hence, the rate of information exchange, typically changes with density and density may be affected by the size of the social group. We investigate models in which each individual may be engaged in one of several tasks. For example, the different tasks could represent alternative foraging locations exploited by an ant colony. An individual's decision about which task to pursue depends both on environmental stimuli and on interactions among individuals. We examine how group size affects the allocation of individuals among the various tasks. Analysis of the models shows the following. (1) Simple interactions among individuals with limited ability to process information can lead to group behaviour that closely approximates the predictions of evolutionary optimality models, (2) Because per capita rates of social interaction may increase with group size, larger groups may be more efficient than smaller ones at tracking a changing environment, (3) Group behaviour is determined both by each individual's interaction with environmental stimuli and by social exchange of information. To keep these processes in balance across a range of group sizes, organisms are predicted to regulate per capita rates of social interaction and (4) Stochastic models show, at least in some cases, that the results described here occur even in small groups of approximately ten individuals.     

Predator–prey interactions and metabolic rates are altered in stable and unstable groups in a social fish

Understanding the determinants and consequences of predation effort, success and prey responses is important since these factors affect the fitness of predators and prey. When predators are also invasive species, the impacts on prey can be particularly far-reaching with ultimate ecosystem-level consequences. However, predators are typically viewed as behaviourally fixed within this interaction and it is unclear how variation in predator social dynamics affects predator–prey interactions. Using the invasive eastern mosquitofish Gambusia holbrooki and a native glass shrimp Paratya australiensis in Australia, we investigated how varying levels of social conflict within predator groups influences predator–prey interactions. By experimentally manipulating group stability of G. holbrooki, we show that rates of social conflict were lower in groups with large size differences, but that routine metabolic rates were higher in groups with large size differences. Predation effort and success did not vary depending on group stability, but in stable groups predation effort by aggressive dominants was greater than subordinates. The anti-predator responses of prey to the stability of predator groups were mixed. While more prey utilized shelters when exposed to stable compared to unstable groups of predators, a greater proportion were sedentary when predator groups were unstable. Overall, this study demonstrates predator group stability is modulated by differences in body size and can influence prey responses. Further, it reveals a hidden metabolic cost of living in stable groups despite reduced overt social conflict. For invasive species management, it is therefore important to consider the behavioural and physiological plasticity of the invasive predators, whose complex social interactions and metabolic demands can modulate patterns of predator–prey interactions.     

Resource distribution and group size in the common langur Semnopithecus entellus in southern India

We investigated the influence of resource abundance and distribution on the group size and composition of the common langur Semnopithecus entellus in the contiguous forests of Bandipur National Park, Mudumalai Wildlife Sanctuary and Nagarahole National Park in southern India. We also explored any additional effect of predator pressure and the risk of take-over on the same attributes. Data on group composition and vegetation were collected from January to May 2006. The size and composition of 94 bisexual groups were obtained. The group size varied from 7 to 40 and the groups included unimale and multimale groups. Thirty-five all-male groups were encountered. Vegetation was sampled from 17 grids of dimension 1 km x 1 km, each containing twelve 25 m x 25 m plots. The list of food species was compiled from previous studies and observations made during the study period. The mean basal area of all the food trees within each plot and its coefficient of variation at the level of the grid were used to represent resource abundance and distribution, respectively. The number of adult females and males within groups were analyzed separately to test for differential effects on age-sex categories. Group size increased as resources became spatially more heterogeneous. The abundance of resources had a negative effect on group size. This study did not find evidence supporting the direct effect of predator presence or of the risk of take-over. Contrary to what were expected, adult males reacted more strongly and predictably to resources than did adult females. The group attributes and their relationship with food resource abundance and distribution differed between two sites in the study area possibly owing to langur subspecies differences.     

The building-up of social relationships: behavioural types, social networks and cooperative breeding in a cichlid

Consistent individual differences in behavioural types may not only cause variation in life-history decisions, but may also affect the choice of social partners and sociality in general. Here, we tested whether and how behavioural type influences the establishment of social ties using the cooperatively breeding cichlid, Neolamprologus pulcher. In a habitat saturation experiment with individuals pre-tested for behavioural type, we first analysed whether behavioural type affected the likelihood of settlement (i.e. social status), group sizes, and the types of dominant and subordinate individuals accepted as group members. Corrected for effects of body size and sex, the behavioural type did not affect settlement. However, bold dominant males only accepted smaller females, and grouped with bold subordinates, while shy dominant males accepted larger females than themselves, and grouped with shy subordinates. Second, we analysed the relationships between behavioural type and the aggressiveness or affiliation social network. Behavioural type significantly affected the number and quality of connections within the two networks. We show that behavioural types affect group composition, social networks and status achieved, in interaction with body size. Thus, the interactions within groups may depend not only on age, size and sex, but also on the behavioural type of the individuals involved.     

Demography, range use, and behavior in black lemurs (Eulemur macaco macaco) at Ampasikely, northwest Madagascar

We studied a black lemur population over a 2-year period (1992-1993) and 8 years later (2000) in a 50-ha secondary forest in northwest Madagascar. All of the animals were marked to investigate population dynamics and seasonal variation in ranging and behavior, and new data on black lemurs were obtained. Our data on demographic characteristics were expanded to include other forest sites and contrasted with those collected in other Eulemur macaco macaco field studies, in relation to human activity and the presence of introduced and cultivated plant species. Density is affected by deforestation and hunting. Group size and home range depend on the composition of the forest and probably food patches. Sex ratio at birth varies according to the number of females per group, a result that fits the local resource competition model. Groups are multimale-multifemale, and adult females form the core of the groups. Reproductive parameters indicate sharply defined seasonal breeding, a high female reproductive rate, and birth synchrony. Changes in group composition reveal male and female juvenile dispersal, male transfer between groups at the time of mating, and adult female transfer and group fission when groups exceed a critical size. At mating and birth, intergroup agonistic encounters occurred at home-range boundaries, and larger groups were dominant over smaller groups. Patterns of intragroup interactions suggest that males compete for access to groups of females during the mating season, and that females may compete for food resources during the birth season. Our study also reports female social dominance and lack of sexual weight dimorphism in this species.