Angela M. Sibbald

Allelomimetic synchronization in Merino sheep

Changes between the inactive (resting/ruminating) and active (grazing, walking) states in groups of Merino sheep were studied in the field for different group sizes (two, four, six or eight) of either male or female animals over 6-h periods. The amount of synchrony within groups was high (60–80%) and is attributed to the mutual adjustment of behaviour by group members. To quantify this process, changes in the number of active individuals were fitted by a time homogeneous continuous time Markov chain model. We found that the probability of an individual becoming active increased with the number of active conspecifics in the group and decreased with the number of inactive conspecifics. The reverse effect was found for the probability of becoming inactive. A model of this individual decision-making process is fitted to the data and predictions of the model are shown to account for the synchrony observed within the group. Group synchronization is thus presented as a self-organized dynamic system, where collective oscillations between activity and inactivity arise stochastically from the coupling between individual Markov processes.

Personality and collective decision-making in foraging herbivores.

The mechanisms by which group-living animals collectively exploit resources, and the role of individuals in group decisions, are central issues for understanding animal distribution patterns. We investigated the extent to which boldness and shyness affect the distribution of social herbivores across vegetation patches, using sheep as a model species. Using an experimental and a theoretical approach, we show that collective choices emerge through the nonlinear dynamics of interactions between individuals, at both short and long distances. Within a range of parameter values derived from the observation of homogeneous groups of each behavioural type, we propose a simple mechanism whereby the same interaction rules can result in different patterns of distribution across patches for bold and shy individuals. We present a mathematical model based on behavioural rules derived from experiments, in which crowding and conspecific attraction affect the probability of entering or leaving patches. Variation in the strength of social attraction is sufficient to account for differences in spatial distribution across patches. The model predicts that resource fragmentation more strongly affects the distribution patterns of shy groups, and suggests that the presence of both bold and shy individuals within groups would result in more flexible behaviour at the population level.

Red deer Cervus elephus vigilance behaviour differs with habitat and type of human disturbance

Abstract Red deer Cervus elaphus, even in wilderness areas, are increasingly exposed to disturbance from human recreation as well as hunting, and it has been suggested that both types of disturbance may be perceived as a predation risk. We studied the vigilance behaviour of red deer in the Scottish Highlands, in sites with traditionally high numbers of visitors (‘disturbed’) and sites with relatively few visitors (‘less-disturbed’) during the main recreational season (spring and summer), and in their mating grounds during the hunting season (autumn and winter). We carried out direct observations, using scan sampling at 3-minute intervals for 1-hour periods, and recorded the number of animals in each group that were vigilant and their mode of vigilance. During the recreational season, in both the disturbed and less-disturbed sites, data were collected in habitats with different levels of cover (grassland, heather and woodland). The percentage of animals that were vigilant was higher in disturbed than in less-disturbed sites, and higher in disturbed grassland (poor cover) and heather (intermediate cover) than in disturbed woodland (good cover). The majority of the vigilant animals in disturbed heather and woodland habitats and in all the less-disturbed habitats were standing. In disturbed grassland, however, lying was the main posture whilst vigilant. In both disturbed grassland and heather, the percentage of vigilant animals that were moving was higher than in woodland or the less-disturbed habitats. In disturbed sites, the deer were more likely to be aggregated when vigilance levels were high. During the hunting season, the overall level of vigilance was higher than at any sites during the recreational season, and the majority of vigilant animals were moving. We conclude that red deer respond to disturbance from human recreational activities by increasing their level of vigilance, but that the nature of their response varies with the level of cover available. We suggest that red deer may lie down when keeping vigil in grasslands, because lying animals are less conspicuous and the low cover will still allow animals to scan their surroundings. We conclude that, although they respond to both types of disturbance by increasing vigilance, red deer perceive human recreation as a less acute threat than hunting.

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.

Moving together: Incidental leaders and naïve followers

Elucidating whether common general mechanisms govern collective movements in a wide range of species is a central issue in the study of social behaviour. In this paper, we describe a new experimental paradigm for studying the dynamic of collective movements. Some sheep (Ovis aries) were first trained to move towards a coloured panel, in response to a sound cue. We present data comparing the behaviour of test groups composed of one of the trained sheep and 3 naïve sheep, and control groups composed of 4 naïve sheep. In the tests, for both test and control groups, sheep were observed for 20 min before the sound cue was delivered and the panel made visible. Before the sound, trained and naïve sheep were similar in terms of activity budgets, spatial distribution, social behaviour and spontaneous movement initiation. After the sound, trained sheep moved toward the panel and systematically triggered a collective movement in all test groups. The results suggest that any individual moving away from the group can elicit a collective movement. Our experimental protocol provides an opportunity to quantify mechanisms involved in group movements, and to investigate differences between species and the effect of social context on collective decision-making.

The influences of nutrition and management on the growth of red deer calves from weaning to 16 months of age

The effects of date of weaning, weaning live weight, autumn grazing management and level and pattern of winter nutrition in relation to summer grazing on the live-weight gain of farmed red deer calves to 16 months of age were tested in three experiments. In experiment 1, the effect of two weaning live weights (37 and 46 kg) in September and two levels of autumn and winter nutrition were examined with 48 calves. In October, live-weight gains were positively related to sward height and herbage mass. The calves lost live weight in November. Compensatory live-weight gains of 28 g/day occurred at summer pasture after the low level of winter nutrition. Weaning live weight had a major influence on live weight at 16 months of age. In experiment 2, the effects of maintenance, ad libitum and intermediate treatments were compared using 50 calves given a complete diet in winter. The intermediate treatment consisted of a maintenance followed by an ad libitum level of feeding. Compensatory live-weight gains of 49 g/day occurred at pasture with the intermediate treatment achieving complete live-weight compensation at 16 months of age. In experiment 3, the effect of two weaning dates (19 September and 14 November) on live-weight gain of calves and on the date of subsequent calving of hinds were compared with 34 lactating hinds. There was no difference in live-weight gain of calves between weaning dates but the subsequent calving date was delayed by 8 days from those hinds weaning in November. A management system is advocated with weaning in September, housing in November after grazing sown swards greater than 6 cm, a pattern of a low followed by a high level of nutrition in winter and the summer grazing of sown swards maintained at 4 to 6 cm.

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.

Effect of dosing regime on intake estimation using the n-alkane technique in sheep fed pelleted grass meal.

An experiment was carried out to evaluate the n-alkane technique for estimating the voluntary intake of pelleted grass meal by sheep. Two groups of 12 Scottish Blackface sheep were dosed once daily or twice daily with paper pellets containing 130 mg of C32 alkane. Mean daily intakes over a 5 day period were estimated from n-alkane concentrations in faeces and compared with actual intakes measured by weighing of food. Two faecal sampling frequencies were compared (once or twice daily) at each dosing frequency. Diurnal patterns of intake were determined by weighing food residues every 3 h for 4 days. Diurnal patterns of faecal n-alkane concentration were determined from rectal grab samples collected every 3 h for 2 days. There was no effect of dosing frequency on actual intake. Estimated intake was higher for twice-daily than once-daily dosed animals when twice-daily faecal sampling was employed (P < 0.05), although differences between mean actual and estimated intakes were not significant. The accuracy of estimation of individual intakes was better for sheep dosed once daily, at both faecal sampling frequencies, than for sheep dosed twice daily. This may have been due to diurnal variation in food intake and digesta outflow from the rumen, demonstrating the importance of matching n-alkane dosing and sampling regimes to the digesta kinetics of the diet fed. © 2000 Society of Chemical Industry

Effects of human disturbance on the diet composition of wild red deer (Cervus elaphus)

Disturbance from human recreational activities may affect the nutrition of free-ranging herbivores due to trade-offs between feeding in preferred habitats and perceived predation risk. To test this hypothesis, we estimated diet composition for red deer in the Scottish highlands in spring, when recreational activity tends to be high, and in winter when it is lower. We analysed faecal samples from three habitat types (grassland, heather moorland and woodland) collected at sites close to a busy track (disturbed) and at a distance from it (less disturbed). The diet consisted of 39% grasses, sedges, herbs and rushes (GSHR) and 58% Calluna vulgaris and Erica spp. (heather) in spring, compared with 14% grasses and 77% heather in winter, with small quantities of Vaccinium spp. (berry) and Pinus sylvestris (tree) in both seasons. In spring, faeces from disturbed grass and woodland sites indicated a diet with less GSHR and more heather and tree than faeces from less-disturbed sites, which could be due to an increased need for vigilance in exposed grassland and the need to seek cover. In contrast, faeces from all disturbed sites in winter indicated a diet with more GSHR and less heather than faeces from less-disturbed sites. This could be due to a seasonal decline in recreation and increase in hunting activity reversing the disturbance levels at the different sites, since hunting is not normally carried out in areas used by the public for recreation. We conclude that there may be nutritional benefits to deer of reducing disturbance near open grassland.