Vigilance during Preening in Coots Fulica atra

In birds, vigilance during feeding is usually linked with (1) the many‐eyes‐hypothesis, (2) the dilution effect, and (3) the intraspecific scramble competition hypothesis. To exclude competition for resources as a driving force in vigilance research into activities other than feeding is necessary. Additionally, variables such as nearest neighbour distance and position within a flock are supposed to influence vigilance and vigilance of singletons should be compared with vigilance of flock members. I studied vigilance during preening in Eurasian Coots (Fulica atra) and counted the number of scans per minute. A total number of 117 coots were sampled with 16 of them preening alone. Coots preening alone showed a significantly higher scan rate. I found a significant negative correlation between vigilance and flock size while nearest neighbour distance correlated positively with vigilance. Further, vigilance was higher in individuals at the edge. A general linear model using nearest neighbour distance and flock size within 10 m as covariates also revealed a significant influence of flock size but not of nearest neighbour distance. The centre‐edge effect still remained significant. These results indicate that flock size is the most influential predictor of vigilance in preening coots, followed by spatial organization. However, using nearest neighbour distance instead of flock size also produced a significant model as did the flock size measurement within a radius of 20 m. As scramble competition could be ruled out, the flock size effect may be indeed related to predation.     

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.     

Exchangeable producer and scrounger roles in a captive flock of feral pigeons: a case for the skill pool effect

We investigated the foraging producer-scrounger system of a captive flock of feral pigeons (Columba livia) by monitoring the number of food patches each individual produced. In one experiment, three different patch types were tested on the whole flock while, in a second, flock composition was varied for one patch type. In all cases we found non-uniform distributions of the number of patches produced per individual, which suggests the existence of producer and scrounger roles. This result could not be explained by either dominance or variability in individual learning ability. Individuals switched roles in response to changes both in food patch type and flock composition. These results are discussed in light of the skill pool hypothesis, which suggests that, in a group, different foraging specialists will profit by parasitizing each other's food discoveries.     

Acoustically mediated individual recognition by a coral reef fish (Pomacentrus partitus)

Acoustically mediated individual recognition has been demonstrated in the field by males of the bicolor damselfish Pomacentrus partitus Poey. Playbacks of non-resident sounds from a given fish's territory elicited a greater response from its nearest neighbour than playbacks of the resident's sound—in accordance with the predictions of the ‘dear-enemy’ effect. Testing also included switching the sounds of the two nearest neighbours relative to each respective male's territory. Results demonstrated that all males in the colony individually recognized the sounds of their two nearest neighbours.     

Effects of group size on school structure and behaviour in yellow‐eyed mullet Aldrichetta forsteri

Schooling behaviour in yellow‐eyed mullet Aldrichetta forsteri, a common fish species in New Zealand estuarine habitats, was investigated to identify interaction rules associated with group formation. Tank‐based three‐dimensional studies of three group sizes (15, 75 and 150 individuals) were carried out to measure the effects of these different group sizes on school structure during control, predation risk and foraging behavioural states. Increased group size positively correlated with nearest‐neighbour distance in control and foraging states. Swimming speed was the lowest in all three behavioural states in groups of 15 fish compared with 75 or 150. Immediate behavioural response following visual exposure to a simulated avian predator differed between groups resulting in loss of structure in larger groups. School shape was an oblong–oblate spheroid with a length, breadth and height ratio of 5:2:1 and the area of free space surrounding individual fish was spherical in shape with a high degree of spatial isotropy present in all size groups. These findings challenge traditional theories based on either local or global properties as key drivers of group structure. Instead, our results suggest that a more collaborative approach involving both group size and rules pertaining to nearest‐neighbour interactions affects collective behaviours in this species.     

Coots Fulica atra reduce their vigilance under increased competition

Birds frequently interrupt feeding to scan their surrounding environment. Usually an inverse correlation between scan rate and flock size exists. The 'many-eyes' hypothesis suggests that more eyes are able to detect a predator earlier. Due to the 'dilution-effect' animals in larger groups experience 'safety in numbers', while the 'scramble competition' hypothesis suggests that individuals reduce their vigilance in larger groups since they compete for a limited amount of scarce resources. Here, I induced competition in natural coot populations. Coots preferred feeding on grass and were occasionally fed by passers-by at the study sites. However, this resource was scarce and coots experienced additional food as a very limited resource. I sampled coots prior and after an experimental treatment with an additional food supply. Correlations between vigilance rate (number of scans during 1 min of foraging), nearest neighbor distance and flock size existed before the experimental treatment but afterwards the correlation between vigilance and nearest neighbor distance faded. I found a significantly lower vigilance rate and nearest neighbor distance after inducing competition. This study provided experimental evidence for the 'scramble competition' hypothesis namely that individuals in groups lowered their vigilance when faced with competition for a limited amount of food resources given similar flock sizes.     

Inter-male Spacing and Aggression in African Painted Reed Frogs,Hyperolius marmoratus

Inter-male spacing and the role of aggression in the maintenance of spatial organization was examined in choruses of Hyperolius marmoratus males housed in a semi-natural enclosure. The effects of chorus size on spacing patterns, nearest neighbour distances and male aggressive behaviour were examined. Males were found to reduce nearest neighbour distances as chorus size increased. This was accompanied by a shift in spacing pattern from even in small choruses (5–9 males) to random in choruses of 10 to 13 males. In choruses of 14, spacing patterns were once again even. The reason for these shifts is unclear but may reflect a sudden, rather than gradual increased tolerance by individuals to high neighbour call intensity as space to call from becomes more limited. The level of male aggression was influenced by chorus size and the time of night. In general, more aggressive interactions occurred in high density choruses. However, this did not translate into higher levels of individual male aggression as density increased. Individual male aggression was high in early evening choruses and declined to a minimum at peak chorusing time. High levels of aggression during early evening may reflect the establishment of calling sites by males, while the drop in aggression at peak chorusing time may occur in response to the presence of females in the chorus at this time or as a consequence of masking of neighbours calls.     

Flock density, social foraging, and scanning: an experiment with starlings

Social foraging differs from individual foraging because italters both resource availability and the forager's behavior.We examined responses of starlings to the presence of conspecificsby manipulating foraging-group density experimentally, whileensuring that each subject's foraging opportunities were unchanged.To do this, we used individuals foraging simultaneously in fourbottomless enclosures placed at various separations in naturalforaging grounds. We measured foraging and scanning intensityand qualitative aspects of scanning of focal individuals. Additionally,we examined the temporal distribution of scanning between individuals.The focal individual analysis showed that (1) food-searchingactivity increased, while time spent scanning, time off theground and scanning bout length decreased with flock density;(2) food finding per unit of searching effort increased withdensity; (3) head orientation during scanning was sensitiveto companions' proximity: heads pointed away from the companionsat close distance, toward them at intermediate distance, andwas random farther away. The analysis of the (temporal overlappingin scanning) temporal distribution of scanning for the groupshowed that scanning was significantly synchronized when companionswere adjacent to each other but was not significantly differentfrom random at further separations. We conclude that behavioralresponses of individuals to the presence of others generateimportant changes in foraging performance even in the absenceof physical interference and, more generally, that assessingthe mechanisms that control the behavior of group members atdifferent flock densities offers a way to understand the functionaland ecological significance of foraging aggregations.     

Vigilance Patterns of Territorial Guanacos (Lama guanicoe): The Role of Reproductive Interests and Predation Risk

We conducted focal observations of territorial guanacos, a highly polygynous and social mammal, to compare time budgets between sexes and test the hypothesis that the differences in reproductive interests are associated with differential group size effects on male and female time allocation patterns. In addition, we used group instantaneous sampling to test the hypothesis that grouping improves detection capacity through increased collective vigilance. We fit GLM to assess how group size and group composition (i.e., presence or absence of calves) affected individual time allocation of males and females, and collective vigilance. As expected from differences in reproductive interests, males in family groups devoted more time to scan the surroundings and less to feeding activities compared to females. Both sexes benefited from grouping by reducing the time invested in vigilance and increased foraging effort, according to predation risk theory, but the factors affecting time allocation differed between males and females. Group size effects were significant when females were at less than five body‐lengths from their nearest neighbour, suggesting that grouping benefits arise when females are close to each other. Female time budgets were also affected by season, topography and vegetation structure. In contrast to our expectation, males reduced the time invested in vigilance as the number of females in the group increased, supporting the predation risk theory rather the intrasexual competition hypothesis. The presence of calves was associated with an increase in male individual vigilance; and vegetation type also affected the intensity of the group size effect over male time allocation. In closed habitats, collective vigilance increased with the number of adults but decreased with the number of calves present. Although male and female guanacos differed in their time allocation patterns, our results support the hypothesis that both sexes perceive significant antipredator benefits of group living.     

Factors causing variation in flock size: Decision making to join a foraging flock

In order to clarify the temporal variation of flock size, we examined factors that cause the temporal variation of foraging flock size in a wintering population of the Yellow-throated bunting (Emberiza elegans). We conducted field observations to examine whether the flock size varies greatly and whether non-random associations between individuals exist. We confirmed that flock size varied greatly and some individuals showed non-random associations with other flock members. Then, we carried out computer simulations that assume stochastic risky situation, the condition of hunger level and non-random associations between individuals. In the simulation, the condition of hunger level caused the variation of flock size. However, the distribution of flock size differed from observed ones. When the condition of non-random association was added to the simulation, the variation of flock size became large and the distribution of flock size was similar to that of the observed one. In wintering flocks of Yellow-throated bunting, each individual attempts to forage in an optimal flock size. However, if they are extremely starved, it is suggested that they adopt a conditional strategy to join a flock independently of the flock size to acquire the energy necessary for survival. Simultaneously, they may decide to forage under the effect of non-random association between individuals.     

Scent marking by common volesMicrotus arvalis in the presence of a same-sex neighbour

Several vole species use scent marked runways radiating from their burrows for foraging and dispersion. These marks are probably used for social communication. This 4-day laboratory study investigated the environmental and social causations of marking inside pre-existing corridors in male and female common volesMicrotus arvalis (Pallas, 1778). Firstly I tested the novelty and the reinforcement hypotheses in isolated voles, predicting respectively a habituation or a continuous increase in mark deposition. I then confronted with each other two same-sex voles for two days to investigate the differences between males and females in the pattern of marks inside three corridors, one of which runs along the common partition with the neighbour. I tested the self-advertisement and territorial-defence hypotheses, respectively predicting in the presence of a neighbour either a similar marking between the three corridors or a greater marking in the corridor close to the neighbour than in the two other corridors. The results showed no habituation in marking, even in a familiar environment, confirming the reinforcement hypothesis. After the addition of a neighbour, only the females left more marks in the corridor that ran alongside the common border than in the two other corridors. The territorial-defence hypothesis was thus confirmed in (territorial) females while the self-advertisement was supported in (non-territorial) males. Finally, I tested the competitive-ability hypothesis in females, stating that the abundance of scent marks of an individual before a social interaction can predict its degree of intolerance in a future social interaction. The results from female pairs physically interacting for four days support the hypothesis.     

Predation risk, gender and the group size effect: does elk vigilance depend upon the behaviour of conspecifics?

Many animals benefit from the presence of conspecifics by reducing their rate of scanning for predators while increasing their time spent foraging. This group size effect could arise from a decreased perception of individual risk (dilution hypothesis) and/or an increased ability to detect predators (detection hypothesis). We compared individual and group vigilance of Rocky Mountain elk, Cervus elaphus, in three regions of Yellowstone National Park, Wyoming, U.S.A. that varied in their encounter frequency with coyote, Canis latrans, grizzly bear, Ursus arctos, and grey wolf,Canis lupus , predators. Adult females without calves increased scanning and decreased foraging with high encounter risk and small herd size. Adult females with calves increased scanning and decreased foraging with high encounter risk, but showed no decrease in scanning with large herd size. Yearlings increased scanning and decreased feeding with small herd size, but not with high encounter risk. Adult males were least vigilant, fed most and were not influenced by encounter risk or herd size. These age-sex class differences led to significant differences in group vigilance depending on the composition of the herd. Herds with a majority of mothers were significantly more vigilant than herds with a majority of adult males. However, these differences in group vigilance had no influence on the individual scanning of females without calves. Thus, the decrease in individual scanning with herd size may depend more on changes in individual risk than on cooperative detection of predators. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Social cohesion in groups of sheep: Effect of activity level, sex composition and group size

We investigated the effects of activity, group size and sex composition on the cohesion of merino sheep (Ovis aries) groups. Mixed-sex (50% of each sex) and single-sex groups of 2, 4, 6 and 8 sheep were placed within 491-m² arenas located in natural pastures and were video recorded during 6 daily hours. The behaviour, orientation and location of each sheep were then extracted from the films at 1-s intervals. We analysed the polarisation of individual orientations, mean inter-individual and nearest neighbours’ distances, as well as the frequency of pairs of nearest neighbours according to their sex within mixed-sex groups. Sheep were more aggregated than predicted under the null hypothesis of random spatial distribution for all group compositions and sizes. Sheep were more spread out and less aligned in half-active than in all-active groups, showing that social cohesion was reduced by a lack of activity synchronisation. The highest proximity between individuals was found in resting groups, yet alignment was low. The polarisation peaked in all-active groups. Mean inter-individual distance did not vary and the nearest neighbour distance decreased as group size increased. When sheep were all-active or all-resting, mixed-sex groups were more spread out than single-sex ones, with a greater distance between opposite than between same-sex individuals. Nearest neighbours of the same sex were also more frequent than random. Our results show that social cohesion can be modulated by activity synchrony but also by social affinity.     

Neighbourhood analysis as an indicator of spatial requirements of broiler chickens

The appropriate stocking density for broiler chickens is a much discussed topic in animal welfare. To determine at which stocking density the level of crowding becomes aversive to 4–6-week-old broiler chickens, spatial distribution and behaviour of groups stocked at 8, 19, 29, 40, 45, 51, 61 or 72 birds per 3.3 m² were analysed. Spatial distribution was evaluated using three different indices: inter-individual distances, nearest neighbour distances and Dirichlet polygon areas. The assumption was that broilers would increase the distance to their pen mates if high densities (i.e., close proximity to pen mates) were experienced as aversive, whereas they would decrease this distance if close proximity was experienced positively. Increased distances to pen mates would lead to increased nearest neighbour distances and a more homogeneous distribution (i.e., lower variation of inter-individual distances and of Dirichlet-polygon size) than expected by chance. The distribution expected by chance was determined from both a random distribution and a ‘resource-corrected’ random distribution (which incorporated environmental influences on spatial distribution but excluded social ones).Behavioural observations showed that at higher stocking densities more sitting bouts (P = 0.003) and adjustments of the sitting and lying posture (P < 0.001) occurred. It was also found that nearest neighbour distance varied according to behaviour (P = 0.001). Birds that were eating/drinking were further apart from their nearest neighbour than birds that were foraging, preening, adjusting their sitting or lying posture or showing “other” behaviour.The results from all three methods of spatial analysis suggested that broilers in groups ≥19 birds per 3.3 m² (ultimately equivalent to 15 kg/m²) started to experience the proximity of conspecifics as aversive at some point during the last 3 weeks of rearing. However, nearest neighbour distance analysis showed evidence of aversiveness earlier in life than the other methods of analyzing spatial distribution (variation in inter-individual distance and polygon size), suggesting that nearest neighbour distance is the more sensitive indicator of space requirements.When uneven use of the different areas within the pen was reflected in the expected distribution (i.e., for comparisons to the resource-corrected random distribution) different results were obtained than when such measures were omitted (i.e., for comparisons to the random distribution). As such, this study emphasises the importance of accounting for environmental influences on distribution within a pen.     

Temporal and spatial patterns in the emigrations of the army ant Dorylus (Anomma) molestus in the montane forest of Mt Kenya

Abstract.  1. The emigration behaviour of the army ant Dorylus ( Anomma ) molestus was studied in the montane forest of Mt Kenya. This species forages by massive swarm raids (mean width 10.3 m ± 4.6 m SD), which are assumed to have a strong negative impact on the densities of prey populations.
2. For non-reproductive colonies the stay duration in a nest is highly variable (median 17, range 3–111 days). This suggests that the frequency of emigrations is not dictated by a brood cycle as an underlying endogenous pattern generator.
3. Colony density is high (mean nearest neighbour's distance 82.1 m ± 29.4 m SD) and mean foraging range is 75.0 m, so encounters with neighbouring colonies occur frequently.
4. The straight line emigration distance is on average 92.7 m (± 29.7 SD). The emigration direction is random with respect to absolute bearing and also relative to the direction of the previous emigration. However, the emigration direction is influenced by the location of the nearest neighbour. Colonies typically emigrate directly away from their nearest neighbour.
5. Local food depletion is likely to be the ultimate cause for emigrations in this species, because emigration distance is larger than foraging range and colonies move away from their nearest neighbour. A small percentage of emigrations may be triggered by pangolin attacks on nests.
6. Contrary to the prediction of a recently developed mathematical model for epigaeic swarm-raiding Dorylus ( Anomma ) species, D.  ( A .)  molestus colonies do not engage in intraspecific battles. Possible reasons for the absence of fights between colonies despite apparently fierce intraspecific competition are discussed.     

Costs and benefits of flocking in foraging white-fronted geese (Anser albifrons): effects of resource depletion

This study investigated the costs and benefits of flocking in white-fronted geese Anser albifrons foraging on rice grains in Japan. The time budgets of focal geese were recorded, and the effects of flock size on the proportions of time spent in vigilant and agonistic behaviour were tested. The results showed that the decline in vigilance level and consequent increase in foraging time were beneficial results of flocking whereas agonistic interactions, a potential cost of flocking, did not increase with increasing flock size. However, seasonal variation in flock size suggested that exploitative competition could be a cost of flocking; the sizes of flocks in spring, when resource depletion had progressed, were significantly reduced compared with those in autumn. An experimental increase in rice density resulted in a significant increase in flock size. We conclude that the flock size of foraging white-fronted geese is a result of compromise between a constant benefit of flocking (i.e. decline in vigilance level) and a cost of flocking varying with food abundance (i.e. exploitative competition).     

Predator avoidance as a function of flocking in the sexually dichromatic Hawaii akepa

Hypotheses for joining a mixed-species bird flock consider each species as a single unit. In sexually dichromatic birds, differential conspicuousness between the sexes may result in differences in vigilance for predators. Aspects of the predator avoidance and foraging enhancement hypotheses for the selective value of joining a mixed-species flock were assessed for the strongly sexually dichromatic Hawaii akepa (Loxops coccineus coccineus). There was support for the primary predictions of the predator avoidance hypothesis: vigilance levels decreased with increasing group size, and with membership in a flock, but only for brightly colored adult males. There was little support for the hypothesis that the primary benefit of joining a mixed-species flock is to enhance foraging efficiency through local enhancement.     

Vigilance in Greater Flamingos Wintering in Southern Tunisia: Age-Dependent Flock Size Effect

Decrease in individual vigilance with flock size is a widely recognized pattern in group‐living species. However such a relationship may be affected by other factors, such as age and flock composition. For instance, because young animals generally lack experience and have higher nutritional needs than adults, they can be expected not only to be less vigilant than adults but also to decrease their vigilance level by a greater extent when flock size increases than adults do. We investigated this issue using data on greater flamingos wintering in the gulf of Gabès, in southern Tunisia. Flamingos tended to congregate in small single‐age flocks for feeding, but as flock size increased, flocks became mixed. We found that when flock size increased, young flamingos significantly decreased their vigilance time, while adult did not, suggesting an age‐dependent flock size effect on vigilance. However, when flock composition (single‐age vs. mixed) was taken into account, a more complex pattern was found. Within single‐age and small flocks, no difference was found between young flamingos and adult ones regarding their vigilance level and their response to increasing flock size. However, within mixed and large flocks, adult flamingos were more vigilant than young ones, while variation in flock size did not result in a significant change in vigilance. These results suggest that young birds relied on the presence of adults, and hence more experienced individuals in detecting dangers, to reduce their vigilance and to increase their foraging time in order to satisfy their higher nutritional requirements. They could also be interpreted as a possible consequence of increasing competition with flock size which constrained more nutritionally stressed young flamingos to increase their foraging time to the detriment of vigilance.     

Effects of size and sex on the courting success and foraging behaviour of Embiotoca jacksoni

To test life-history theory that body size and sex should influence how animals allocate time to foraging versus reproductive activities, we measured the effects of size and sex on courting success and foraging behaviour of black surfperch Embiotoca jacksoni off Santa Catalina Island, southern California. Observations of focal fish were made while snorkelling, during which the length of each fish (estimated to the nearest cm), total duration of courting encounters and foraging rates were recorded. We made observations during and outside the mating season. Courtship occurred only between pairs and its duration increased with the size of both the male and female. Although males would court females that were smaller or larger than themselves, pairs that were closely matched in size had long courting sessions, whereas those that differed considerably in size courted only briefly. Small fish foraged more than larger fish, both during and outside the mating season. Males and females foraged at similar rates outside of the mating season, but during the mating season males reduced their foraging rates to less than half that seen outside of the mating season, whereas females continued to forage at the same rate. This decrease in foraging rate of males during the mating season was seen in all sizes of males but was proportionally greatest in the largest males. These observations indicate that males trade off time spent on foraging for time spent courting during the mating season, whereas females do not.     

Spatial pattern,regeneration and growth rates of Brachystegia spiciformis and Julbernardia globiflora

Past work on tree-to-tree interactions in semi-arid savannas, through the use of nearest neighbour analysis, has shown both a regular dispersion pattern and a positive correlation between the size of a tree and the distance to its nearest neighbour. From these results, the importance of competition has been inferred. In the present study, tree-to-tree interactions in a mesic savanna woodland at Marondera, Zimbabwe, dominated by relatively small-sized individuals of Brachystegia spiciformis and Julbernardia globiflora, were examined. Results show that the regular dispersion pattern is uncommon; that there are few significant positive correlations between size of individual and distance to nearest neighbour; and that growth rate is not consistently correlated with size and distance of nearest neighbour. Results of a study of regeneration relative to canopy cover suggest a possible mechanism for these findings: unlike semi-arid savannas, young plants often grow in under-canopy environments so that early spacing of young trees does not occur. Significant positive correlations between the size of a tree and the distance to its nearest neighbour were found in a stand of larger, presumably older, trees, suggesting that such a relationship only develops in mesic miombo woodland through a thinning process as the trees mature. Familial clumping, which is common in the study area, may mask spatial patterns.Abbreviations RBAI Relative basal area increment