Sexual dimorphism, activity budget and synchrony in groups of sheep

The activity budget hypothesis has been proposed to explain the social segregation commonly observed in ungulate populations. This hypothesis suggests that differences in body size – i.e. between dimorphic males and females – may account for differences in activity budget. In particular, if females spend more time grazing and less time resting than males, activity synchrony would be reduced. Increased costs of maintaining synchrony despite differences in activity budget would facilitate group fragmentation and instability of mixed-sex groups. In this paper two prerequisites of the activity budget hypothesis were tested: (1) that males should spend less time feeding and more time resting than females in single-sex groups and (2) that lower activity synchrony should be observed in mixed-sex compared to single-sex groups. The activity budget and synchrony in mixed and single-sex groups of merino sheep (Ovis aries) of different sizes (2, 4, 6, 8 individuals) were measured in three contiguous 491-m² arenas located in a natural pasture. Three same-size groups, one of each category, were observed simultaneously. We found no sexual differences in the time spent inactive and active (i.e. grazing, standing, moving, interacting). Males spent significantly more time grazing and less time standing than females. These differences disappeared when yearling males were omitted from the group. Males and females had similar bite and step rates. Sheep of both sexes spent less time resting and more time grazing and moving and had lower bite rates when in mixed-sex groups than when in single-sex groups. The synchrony among visually isolated groups was near zero, indicating that they changed activities independently. On the contrary, within-group synchrony was high; however it was higher in single-sex groups, in particular for males, than in mixed-sex groups. Our results suggest that differences in activity budget and synchrony alone are insufficient to explain social segregation.     

Could asynchrony in activity between the sexes cause intersexual social segregation in ruminants?

In many sexually dimorphic mammal species, the sexes live outside the mating season in separate social groups (''social segregation''). Social segregation occurs in a wide range of environmental conditions, but its cause in unknown. I suggest that social segregation is caused by a lower level of activity synchrony between individuals in mixed-sex groups than in single-sex groups, owing to sex differences in activity rhythm. As a consequence, mixed-sex groups are more likely to break up than single-sex groups, resulting in a predominance of single-sex groups at equilibrium. To test this hypothesis in red deer (Cervus elaphus L.), I developed an index of activity synchronization and showed that deer in mixed-sex groups were significantly less synchronized in their activity than deer in single-sex groups. Thus, low intersexual synchrony in activity can lead to social segregation. However, a lower level of intrasexual (female-female and male-male) activity synchrony within mixed-sex than within single-sex groups implies that additional factors (other than sex differences in foraging rhythm) contribute to the higher degree of instability of mixed-sex groups.     

Sexual segregation in fallow deer: are mixed-sex groups especially unstable because of asynchrony between the sexes?

In gregarious ruminants, females and males tend to live in separate groups outside the rutting season. According to the 'activity budget' hypothesis, this is due to an activity asynchrony between the two sexes reducing the lifetime of mixed-sex groups. We tested this hypothesis in a fallow deer population. Activity asynchrony was more frequent in mixed-sex than in single-sex groups. In addition, mixed-sex groups had a higher probability of splitting-up than all-female groups, and they mainly split up into single-sex groups. However, activity asynchrony did not appear as a major cause of splitting-up.     

An analysis of factors influencing spatial distribution in flocks of grazing sheep

The spatial distribution of small groups of sheep (5–40) when grazing at a range of densities (4–29 sheep ha^?1) was investigated experimentally. Under these conditions, where the sheep groups rarely split into distinct sub-groups, the angular distributions of the sheep were usually random, but the radial distributions were not. A “concentration curve” representing the density of the sheep as a function of their distance from the centroid of the flock exhibited a peak at a distance varying with breed, flock size and density. Dorset Horn sheep were distributed more widely than Merino, Polwarth, Corriedale or Southdown sheep. Distribution, also measured as area “occupied” per sheep, ranged from 15 to 67 m² among these breeds.Two breeds were studied in detail. Both Merino and Corriedale sheep spread further apart as flock size increased. As density decreased, Merinos but not Corriedales came closer together. There was little correlation in these studies between area occupied per sheep and distance to first nearest neighbour, because distance to nearest neighbour varied relatively little compared with that of average area occupied. Thus, spatial dispersion seems to have two components: (a) a relationship to a neighbour by each individual; (b) spatial cohesion of the group as a whole which is influenced by group size and density.     

Tendency-distance models of social cohesion in animal groups

Although it has been assumed that attraction and repulsion between social individuals constitute a basis for group cohesion, there has been no systematic study of the possible ways in which these tendencies might vary with inter-individual distance (IID), or of associated implications for group structure. In this paper, a family of attraction/repulsion--distance functions is described. Computer simulation was used to examine the effects of each function on group cohesion, as reflected by mean values and variability in IID and group shape. Our results showed that: (a) all models led to stability in group structure, but differed significantly in terms of stable IID and group shape characteristics; (b) cohesion was best served by an upwardly convex behaviour--distance function in which maximum attraction equaled maximum repulsion (and the biological plausibility of this function is discussed); (c) group elongation and variability in mean IID were significantly positively correlated; (d) although dyads maintained an equilibrial separation distance, at which attraction balanced repulsion, in larger groups stable nearest neighbour distances were often less than the equilibrium distance; and (e) individuals needed to monitor and respond to only relatively few of their companions in order to avoid group fragmentation.     

Boys will be boys: social affinity among males drives social segregation in western grey kangaroos

In many sexually dimorphic and polygynous species, individuals exhibit social segregation by grouping with others of their own sex. Several proximate mechanisms have been proposed to explain social segregation: female avoidance of males, male avoidance of females, avoidance of harassment, male social affinity, female social affinity and activity budget asynchrony. Some of these have been tested in ungulate species, but few previous studies have been able to rigorously test, or distinguish between, these mechanisms because they have failed to examine the fission/fusion dynamics of groups. We tested these proximate mechanisms simultaneously in western grey kangaroos Macropus fuliginosus , by examining whether females, small males or large males instigated segregation by leaving mixed-sex groups or joining individuals of their own sex or size class. Females, small males and large males left mixed-sex groups as if leaving were independent of sex–size class. In contrast, large males joined male-only groups more frequently than expected. These results suggest that the need for males to maintain contact with other males can contribute to the cohesion of male-only groups and promote segregation. As male–male competition occurs in many polygynous species that sexually segregate, a comparable mechanism might be operating in other taxa, and should be examined further.     

个体大小对草鱼耐高温能力及升温过程中群体行为的影响

为考察个体大小对升温过程中草鱼的耐高温能力和群体行为的影响, 研究以草鱼(Ctenopharyngodon idellus)幼鱼为实验对象, 结果显示: 大个体群体的个体游泳速度明显大于小个体群体(P=0.039), 个体间距离和最近邻距离显著高于小个体群体(P<0.001); 大个体鱼群的极性与小个体群体并无差异。虽然大个体群体的致死高温与小个体群体无差异, 但前者的临界高温显著大于小个体群体(P<0.001)。升温过程会使鱼群个体游泳速度变快, 个体间距离和最近邻距离也相应增加, 但游泳速度同步性和群体极性维持不变。研究表明: 个体大小的增加可增强草鱼幼鱼对环境水温的耐高温能力; 升温不会影响不同个体大小草鱼幼鱼群体的协调性, 但会降低草鱼幼鱼的群体凝聚力。     

Predation risk sensitivity and the spatial organization of primate groups: A case study using GIS in lowland Woolly Monkeys (Lagothrix lagotricha poeppigii)

Predation risk is thought to be a potent force influencing intragroup cohesion, and the level of risk experienced by an individual is expected to vary with both group size and spatial position within a group. Smaller‐bodied and less‐experienced individuals are presumed to be more vulnerable to predators, suggesting that within‐group spatial organization should show size‐ and age‐dependent patterns in predator sensitive positioning. However, such effects have been difficult to evaluate for arboreal primates living in large groups. We conducted a preliminary study using a novel, spatially explicit method of assessing group spatial organization using GIS data in two groups of wild lowland Woolly monkeys, in which one group had a membership roughly twice as large as the second. In the larger group, group spread was more diffuse and large adult males were more frequently on the outskirts of the group than other age/sex classes, while immatures and females with dependents were more often in the center. Leaf cover around an individual—presumed to index an animal's perception of risk—increased significantly with distance from the group center for all immatures, although they were typically under lower leaf cover than adults; the number of groupmates in proximity also had an effect, but nearest neighbor distance did not. These differences were not detectable in the smaller group. This preliminary study suggests that thorough studies of spatial organization and predation risk sensitivity in arboreal primates are possible and could yield valuable information on how gregarious individuals offset ecological risks through social spacing. Am J Phys Anthropol 156:158–165, 2015 © 2014 Wiley Periodicals, Inc.     

Trade-offs between social behaviour and foraging by sheep in heterogeneous pastures

In heterogeneous pastures, groups of sheep may have to alter their social behaviour in order to graze patches of preferred vegetation. In this event, patch size, inter-patch distance and the contrast between patch and background vegetation are likely to affect behaviour. In this experiment, groups of five female Scottish Blackface sheep grazed for 2-h periods in 0.1ha grass plots containing seven 1.5mx1.5m patches of improved vegetation, with inter-patch distances of 1.5, 6 or 11.5m. Background vegetation was of either medium or poor quality. Control plots contained no patches. On average, sheep spent 44% of the time grazing patches, although patches comprised only 1.6% of the total plot area. Inter-patch distance did not affect accumulated time spent grazing patches during the first 30min, but patch residence time and the number of sheep on a patch increased with inter-patch distance. The distribution of nearest neighbour distances was altered when patches were 6 or 11.5m apart, compared to no patches. Accumulated time spent grazing patches and the number of sheep on a patch were greater with poor than medium backgrounds. Sheep visited patches frequently and for short periods and it is suggested that sheep often moved off patches as a result of competition. The results provide evidence that sheep make trade-offs between social and foraging behaviour and demonstrate the importance of interactions between social spacing and the size and spacing of vegetation patches, when sheep forage in heterogeneous pastures.     

Sex differences in behaviour of the mongolian gerbil (Meriones unguiculatus) in encounters between pairs of same or opposite sex

Pairs of gerbils were observed in mixed-sex and single-sex encounters in a neutral arena using a round-robin technique: various acts and postures were described and their frequency analysed. In single-sex encounters' males showed more social investigation, footstomping and marking while females engaged more frequently in jumping, grooming and chewing. Both sexes showed comparable levels of agonistic behaviour. In mixed-sex encounters, agonistic behaviour was markedly increased due to the aggressiveness of the females, usually in response to sexual investigation by the male. Males never attacked females. High levels of footstomping and marking with the ventral scent gland were characteristic of males in the presence of unreceptive females. Lordosis was only observed in three out of one hundred tests.     

The Role of Neighbours Selection on Cohesion and Order of Swarms

We introduce a multi-agent model for exploring how selection of neighbours determines some aspects of order and cohesion in swarms. The model algorithm states that every agents'' motion seeks for an optimal distance from the nearest topological neighbour encompassed in a limited attention field. Despite the great simplicity of the implementation, varying the amplitude of the attention landscape, swarms pass from cohesive and regular structures towards fragmented and irregular configurations. Interestingly, this movement rule is an ideal candidate for implementing the selfish herd hypothesis which explains aggregation of alarmed group of social animals.     

The number of farm mates influences social and maintenance behaviours of Japanese Black cows in a communal pasture

The objective of the present study was to investigate optimal group size in cattle in communal pastures where cattle from various farms are turned out: (1) to ascertain if cows derived from each farm form an affiliative group and (2) to investigate the effect of the number of farm mates on social and maintenance behaviours when farm mates form an affiliative group. Farm mates were defined as cows from the same farm. A total of 27 cows having zero, one, two to four, or sixteen farm mates were selected as focal animals in two communal pastures. Each focal animal was followed by an individual observer for 8 h from sunrise. The nearest neighbour and the distance to the nearest neighbour were recorded at 10-min intervals, and maintenance behaviours at 1-min intervals; social behaviours and participants were recorded continuously. Most focal animals frequently choose cows from the same farm as their nearest neighbours allogroomed with these farm mates significantly more (P<0.001). Two focal animals did not form affiliative groups with farm mates. The mean distance to the nearest neighbours, whether farm mates or non-farm mates from focal animals having one and two to four farm mates was shorter than cows turned out with no farm mate (1.4 vs. 1.9 times as long as a cow's body length, P<0.05). Cows having two to four farm mates performed (7.8 s/h/cow, P=0.06) and received allogrooming more (9.8 s/h/cow) and escaped less during agonistic encounters (0.1 frequencies/h/cow, P<0.01) than ones having other numbers of farm mates. The mean duration of grazing behaviour per bout tended to increase with the number of farm mates (5.2 to 10.3 min, P<0.05). The mean duration of recumbency behaviour of cows having two to four farm mates (44.0 min/bout) was longer than the ones having other numbers of farm mates. Social and maintenance behaviours of cows having 16 farm mates were like those having zero or one farm mates, which suggested that the social bond among them was weak. It is concluded that a group of cows having two to four farm mates, that is, a group size of three to five cows, may be optimal for a stable life in a communal pasture.     

Some geometric aspects of house sparrow flocks

We analysed 50 movie films of house sparrows (Passer domesticus) foraging on an experimental grid. The location and orientation of each bird was recorded; sex of the individual was determined in about two of every three cases. Results revealed the following: (1) Flock size exhibited a weak inverse relationship to ambient temperature, though aggression was rare in the experimental patch. (2) The frequency of males in foraging groups exceeded the frequency of males in the local population. (3) Within a given flock size, nearest neighbour distances did not differ significantly between male-male and male-female pairs. However, average nearest neighbour distance was inversely related to flock size. (4) Solitaries oriented away from safety and toward a source of disturbance. Orientation of an individual within a larger group was more variable than that of a solitary, and the orientation of nearest neighbours indicated a significant tendency to keep each other in view.     

Black capuchin monkeys dynamically adjust group spread throughout the day

Variation in group spread can affect communication between individuals, synchrony of activity and collective decision-making. Most studies have measured this variation in large time scales, but the variables that influence group spread change as the social unit ranges over different areas throughout the day. Thus, our aim was to analyze the diurnal variation in group spread of a group of black capuchin monkeys (Sapajus nigritus) inhabiting an area within the southeastern Brazilian Atlantic Forest, the Carlos Botelho State Park. We measured group spread as the distance between the coordinates of the location of the two most distant group members every 5 min during entire days. Using time series analysis, we found that the group spread varied within short periods of time (15 min) throughout the day, revolving around the average group spread distance of 36 m. Group members tended to decrease the interindividual distance over time when they were more than 36 m apart, and to increase the distance when they were at a distance of less than 36 m. In addition, group spread was higher when monkeys were eating fruit than during other activities. We conclude that the variation in group spread may reflect competition for discrete food resources and that increasing cohesion several times throughout the day can be a way of resuming social contact between individuals.

     

Effects of space allowance on the grazing behaviour and spacing of sheep

In each of two experiments, four groups of 10 female Scottish Blackface sheep grazed for 2 weeks at each of four space allowances, according to a Greco-Latin Square design. Space allowances were 80, 100, 133 or 200m(2) per head in Experiment 1 and 50, 67, 100 or 200m(2) per head in Experiment 2. Confounding of space allowance and grazing pressure was avoided by the use of a rotational grazing system. Mean distances between sheep were measured from video recordings and distances expected by chance were calculated from simulations in which sheep were located within plot areas using randomly selected co-ordinates.Total time spent grazing was higher at 200m(2) per head and less at 50m(2) per head than at other space allowances, but there was no evidence for an effect of space allowance on herbage intake or digestibility, biting rate or the proportion of grazing time spent in ingestive behaviour. At space allowances from 50 to 133m(2) per head, there were no significant differences between mean observed inter-animal distances and those expected by chance, but observed values were lower than expected values at 200m(2) per head. At all space allowances, mean observed distances between sheep and their nearest neighbours were lower than distances expected by chance.The results show that, on relatively high quality, homogeneous vegetation, female Scottish Blackface sheep will graze at space allowances between 50 and 200m(2) per animal with no evidence of repulsion between individuals and with no clear evidence of grouping until space allowance reaches 200m(2). It is concluded that the relationship between grazing time and space allowance may reflect the extra activity required to maintain group cohesion as space allowance increases.     

Activity synchrony and social cohesion: a fission-fusion model.

A social group can only be spatially coherent if its members synchronize activities such as foraging and resting. However, activity synchronization is costly to individuals if it requires them to postpone an activity that would be personally more profitable in order to do what the rest of the group is doing. Such costs will be particularly high in groups whose members belong to different age, size or sex classes since the optimal allocation of time to various activities is likely to differ between such classes. Thus, differences in the costs of activity synchronization between and within classes could cause non-homogenous groups to be less stable than homogenous groups, with the result that homogenous groups predominate in the population: that is, they could cause 'social segregation' of animals of different sex, size or age. We develop a model that predicts the degree of social segregation attributable to differences in activity synchronization between homogenous and non-homogenous groups and use this model in determining whether activity synchronization can explain intersexual social segregation in red deer (Cervus elaphus). Differences in activity synchronization between mixed-sex and unisex groups of red deer explained 35% of the observed degree of intersexual social segregation, showing that activity synchronization is an important cause of social segregation in this species.     

The formation of preferential relationships at early age in cattle

Calves can develop long-lasting social relationships with peers. We examined the strength of the relationships between calves according to the time they had been together. Twenty-four female dairy calves were assigned to six groups of four animals (Type-1 partners) at 0.5 month of age. At 3.5 months of age, they were mixed with other calves (Type-2 partners) to form groups of 14. Type-3 partners were calves added to the experimental groups after 5.25 months. The calves stayed together until 1.5 years of age. Social preferences between the three partner types were examined in a Y-maze, and the position and activity of animals in the barn and pasture were followed in three periods. Behavioural synchrony, distance between animals, proximity and nearest neighbour were analysed. The calves more frequently butted Type-3 than Type-1 partners in the Y-maze (P < 0.05). They spent more time in proximity to Type-1 partners, and these were more often the nearest neighbours than other partners (P < 0.001). Synchrony and distance between animals were greater at pasture than in the barn (P < 0.01). Calves seem to form preferential relationships before 3.5 months of age. Keeping cattle together from an early age seems beneficial for them.     

Sex differences in the play behavior of immature spotted hyenas, Crocuta crocuta

Social, locomotor, and object play were studied in a colony of five male and five female peer-reared spotted hyenas during 12 1-hr tests while the animals were 13-19 months of age. Animals were tested in both same-sex and mixed-sex groups and were stimulated to play by the introduction of fresh straw and sawdust bedding. Each test was videotaped and the frequency of each type of play was determined by a time sampling procedure. Females played more frequently than males, however, the category of play which was elevated depended upon the social context during testing. In same-sex tests the frequency of vigorous social play displayed by females markedly exceeded that by males, but no comparable sex difference appeared in mixed-sex tests. Females engaged in locomotor play more frequently than males in mixed-sex tests, but no comparable sex difference appeared in same-sex tests. No sex difference in object play was observed. Two male and two female hyenas were gonadectomized prior to the initiation of the present sequence of tests. The results suggest that gonadectomy during the prepubertal period does not affect the frequency of play behavior. However, the small sample sizes preclude any conclusive determination of the effects of these gonadectomies on play.     

Local Mating,Dispersal and Sex Ratio in a Gregarious Parasitoid Wasp

Parasitoid sex ratios can be greatly influenced by mating and dispersal behaviour. Many sex ratio models assume that mating is strictly local (only mated females disperse from the natal patch) and that a single male is sufficient to inseminate all females in a brood. Bethylids (aculeate parasitoids) have been used to test predictions of these models, but less attention has been paid to testing their underlying assumptions. We investigated the timing of eclosion, mating and dispersal in mixed-sex and single-sex broods of the bethylid wasp Goniozus nephantidis. In mixed-sex broods, almost all females mate before dispersal and a single male is sufficient to inseminate virtually all females, even when brood sizes are large. Males disperse from both mixed-sex and all-male broods, but males in all-male broods disperse more slowly. Virgin females disperse from all-female broods, which are common. Virgin females can produce a brood, mate with their own sons and subsequently produce mixed-sex broods, but their success rate is very low. Virgin females could potentially circumvent sex allocation constraints by superparasitizing mixed-sex broods, but when presented with hosts bearing mixed-sex broods they destroy all members of the initial brood before ovipositing. Because of the high prevalence of single-sex broods and dispersal of both sexes, the mating structure of G. nephantidis is unlikely to conform to the assumption of strict local mating.