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.     

Observational and Experimental Evidence for the Function of Tail Flicking in Eurasian Moorhen Gallinula chloropus

Tail movements such as wagging, flicking or pumping are reported from many bird species but their adaptive functions remain poorly understood. To investigate whether tail flicking functions as an alarm signal, either to predators or neighbouring birds, or as a signal of submission to conspecifics, I observed this behaviour in moorhen in a natural context, and conducted playback experiments using vocalizations of predators, conspecifics and heterospecifics. I found positive relationships between flicking and vigilance and nearest neighbour distance, and negative relationships between flicking and moorhen flock size and total flock size. Moorhen at the edge of a flock flicked at a higher rate. Single moorhen flicked more often compared with individuals in groups, both in single‐ and mixed‐species flocks, and there was a tendency that single moorhen flicked more often than single moorhen within a mixed‐species flock. Moorhen responded differently to conspecific and predator calls. While in both cases vigilance increased, tail flick rate was higher during predator playbacks and lower during conspecific playbacks. Furthermore, moorhen remained rather motionless when conspecific calls were played back, but not during predator calls, and, moorhen resumed to a baseline level of tail flicking more quickly after the playback of conspecific calls. Taken together, the results suggest that flicking may be considered as a honest signal of vigilance directed towards ambushing predators.     

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.     

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.     

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.     

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.     

Disturbances by dog barking increase vigilance in coots Fulica atra

Animals frequently interrupt their activity to look up and to scan their surrounding environment for potential predators (vigilance). As vigilance and other activities are often mutually exclusive, such behaviours are at the expense of feeding, sleeping or preening. Authors of many wildlife disturbance studies found that people with free-running dogs provoked the most pronounced disturbances (e.g. greater flushing distances and more birds affected). However, dogs on leash may also negatively affect wild animals, and barking dogs may lead to an increase in vigilance. In this study, I tested this hypothesis in coots (Fulica atra) using three different playback procedures: (1) dog barks, (2) conspecific coot alarm calls and (3) chaffinch song. The trials were conducted in spring and autumn 2005 at three study sites in southwestern Germany. During the dog playbacks, vigilance increased significantly from 17 to 28%. This increase in vigilance is comparable to the presence of a natural predator. As expected, vigilance also increased significantly during conspecific coot alarm calls but not during playbacks of the chaffinch song control. Two main findings result from the study: (1) coots respond to acoustic traits of dogs and may be able to acoustically recognise this predator and (2) this increase in vigilance might have implications for conservation, especially when considering buffer zones around sensitive areas.     

Effects of position and flock size on vigilance and foraging behaviour of the scaled dove Columbina squammata

Amongst the benefits of foraging in flocks are the enhancement of food finding and predation avoidance. Characteristics such as size, individual position, as well as position and distance between members are factors that may influence vigilance and foraging. In a study using scaled doves, Columbina squammata, I observed a negative correlation between group size and vigilance and a positive correlation with time spent foraging, which suggests a reduction of costs and an increase of benefits as a consequence of larger group sizes. Individual position in the flock appeared to be an important factor in this trade-off. Peripheral individuals were more vigilant and foraged less than central ones, suggesting an edge effect in flocks of this species. The clustering of conspecifics may be related with fast transmission of information. Overall, aggressive interactions were rarely observed; when registered, they occurred mostly in larger groups, suggesting an effect of interference competition. These results imply that predation may be a strong pressure on the scaled dove's flock formation and behaviour.     

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.     

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     

The effects of flock size and human presence on vigilance and feeding behavior in the Eurasian Coot (<Emphasis Type="Italic">Fulica atra</Emphasis> L.) during breeding season

Group size influences individual vigilance in different bird species. Most studies have shown that animals spend more time for feeding and less time on vigilance when in groups. Apart from group size, additional factors, such as group density, age, sex, and season, are considered to influence vigilance behaviors. The aim of the present study was to investigate the effects of flock size and human presence on vigilance and feeding in the Eurasian Coot (Fulica atra) during the breeding season. This work was performed in Yorukkirka Lake (Eskisehir, Turkey) from March to August 2005. No correlation was found between flock size and vigilance rates during either human absence or presence. On the other hand, when comparing behavioral differences between human presence and absence, humans appeared to have an important negative effect on vigilance rates, vigilance duration, and feeding behavior, regardless of flock size.     

Temporary intermissions in capturing prey (<Emphasis Type="Italic">Daphnia</Emphasis>) by planktivorous fish (<Emphasis Type="Italic">Rutilus</Emphasis><Emphasis Type="Italic">rutilus</Emphasis>): Are they due to scramble competition or the need for antipredation vigilance?

Capture rates in planktivorous fish may differ in individuals foraging alone or in a group, and this may result either from the altered risk of predation due to vigilance sharing in the group, or from a difference in the intensity of scramble competition for encountered prey items. Changes in capture frequency and the feeding pattern observed in young roach (Rutilus rutilus) feeding alone and in a group of three on a high density prey (Daphnia), in the presence and in the absence of predator odor, were used to determine which of these two alternate explanations is more likely. Earlier studies revealed that a foraging roach captures Daphnia prey in uninterrupted sequences of captures occurring every 1–3 s. Such multiple captures are separated by intermissions of 10–20 s, with their duration being likely to determine the overall capture rate. An experiment was performed to examine whether feeding in a group of three permits higher capture rates (hypothesis 1), and whether the intermittent foraging pattern is due to the need to invest more time for vigilance when foraging alone (hypothesis 2). Video recordings were made of many series of subsequent prey captures by roach feeding on high Daphnia densities, alone or in a group, and in the presence or absence of predator odor. Analysis of these data revealed that the mean duration of intermissions between bursts of feeding activity was significantly greater in the presence of predator odor, which resulted in a significant decrease in the capture rate. Furthermore, when the roach were feeding in a group, these intermissions were reduced to a greater extent in the presence of predator odor than in its absence, implying that the intermission intervals represent an investment for vigilance as an effective antipredation defense that permits increased food intake regardless of whether or not it is enhanced by the resource or the interference competition.     

Sleeping gulls monitor the vigilance behaviour of their neighbours

Individuals in groups are often thought to scan their surroundings for threats independently of one another. Models, however, suggest that foragers should monitor the vigilance level of their neighbours to prevent cheating, and to gather information about incipient predation risk. Evidence for monitoring of vigilance is scant. Here, I examined changes in vigilance levels in sleeping gulls (Larus sp.) surrounded by neighbours in various states of alertness. Controlling for group size and neighbour density, gulls interrupted sleep more often to scan their surroundings, and were therefore more vigilant, when their neighbours were alert rather than sleeping or preening. The results provide evidence for copying of vigilance within groups of birds, suggesting a complex flow of information about predation risk in groups.     

How do vigilance and feeding by common cranes Grus grus depend on age,habitat, and flock size?

Animals often spend less time vigilant and more time feeding when foraging in larger groups. This group-size effect does not, however, consider if larger groups differ systematically from smaller ones: Large groups could form in different habitats than small groups or be composed of a different mix of ages or classes than small groups. We examined how habitat differences and flock size and composition explain feeding and vigilance rates in common cranes Grus grus , wintering in holm oak Quercus ilex dehesas of Spain. Flock size and composition were related to habitat type in cranes: flocks formed in areas sown with cereal crops were larger than flocks formed in set aside areas. Vigilance rate depended on habitat but decreased with increasing flock size in a similar way across all habitats. Juveniles were less vigilant than adults and showed little change in vigilance with flock size. Vigilance increased and feeding time decreased over months from November through February. Our results show that vigilance is affected by habitat but that the group size effect on vigilance is not the product of differences between habitats in group size or composition.     

Effect of Group Size on Individual and Collective Vigilance in Greater Rheas

We studied the effect of group size on the proportion of time that greater rheas, Rhea americana, allocated to vigilance and feeding during the non‐breeding season. We tested whether: (1) the proportion of time that one bird allocates to vigilance (individual vigilance) decreases with group size, and (2) the proportion of time that at least one bird of the group is vigilant (collective vigilance) increases with group size. We analyzed video‐recordings of birds that were foraging alone or in groups from two to 12 birds. The proportion of time allocated to individual vigilance decreased and the proportion of time spent feeding increased with group size. In both cases the main significant difference was between birds foraging alone and in groups. Collective vigilance did not vary with group size and it was lower than expected if vigilance bouts were random or sequential. Our results indicate that rheas foraging in large groups would not receive the benefit of an increase in collective vigilance, although they could still benefit from a reduction of predation risk by the dilution effect.     

Contact calling behaviour in the male ring‐tailed lemur (Lemur catta)

The ring‐tailed lemur (Lemur catta) is a group‐living strepsirhine primate endemic to Madagascar that has a complex vocal repertoire including affiliative moan and hmm calls. Past research has suggested that both vocalisations may be contact calls, and we assessed their usage in addition to examining the effect of individual differences and social factors on male calling behaviour. We tested three hypotheses: the group cohesion, preferred companion and individual differences hypotheses. From March to July 2010, 565 h of focal animal data were collected on 31 males aged ≥1 year at the Beza Mahafaly Special Reserve, Madagascar. Focal males were observed for 30 min, with vocalisations, allogrooming and agonistic interactions noted using one‐zero sampling, and general behaviour, nearest neighbour identity and distance noted using instantaneous point sampling, all at 2.5 min intervals. We found support for the group cohesion hypothesis for both vocalisations, and support for the preferred companion and individual differences hypotheses for the hmm but not the moan vocalisation. Moan and hmm calls maintained the distance between a male and his nearest neighbour, and were made at higher rates during behaviours which may lead to separation from conspecifics, such as travel and vigilance. Males made hmm calls at higher rates when they were low‐ranking, and/or when the nearest neighbour of a male or a preferred companion of either sex. Low‐ranking males may hmm call to maintain proximity to specific individuals within social groups who will show them increased tolerance, with the hmm signalling peaceful intent. Ring‐tailed lemurs are one of the best living models of gregarious primate ancestors, and gaining a better knowledge of the function and range of usage of their contact calls informs our understanding of the evolution of primate cognition.     

Flock feeding and time budgets in the house sparrow (Passer domesticus L.)

A winter population of house sparrows at a farm fed on barley seed in two distinct types of habitat: cattlesheds and open fields. The risk of predation was apparently higher in the fields where birds scanned more frequently than in the cattlesheds and where scanning was negatively influenced by flock size but positively influenced by distance from cover. Individual time budgets were more influenced by flock size than by seed density in the fields but more influenced by seed density than by flock size in the cattlesheds. Higher rates of scanning resulted in greater flock vigilance and longer flight distances in the fields but flight distance was negatively influenced by the density of seeds on which birds were feeding.     

Evolution of local adaptations in dispersal strategies

The optimal probability and distance of dispersal largely depend on the risk to end up in unsuitable habitat. This risk is highest close to the habitat's edge and consequently, optimal dispersal probability and distance should decline towards the habitat's border. This selection should lead to the emergence of spatial gradients in dispersal strategies. However, gene flow caused by dispersal itself is counteracting local adaptation. Using an individual based model we investigate the evolution of local adaptations of dispersal probability and distance within a single, circular, habitat patch. We compare evolved dispersal probabilities and distances for six different dispersal kernels (two negative exponential kernels, two skewed kernels, nearest neighbour dispersal and global dispersal) in patches of different size. For all kernels a positive correlation between patch size and dispersal probability emerges. However, a minimum patch size is necessary to allow for local adaptation of dispersal strategies within patches. Beyond this minimum patch area the difference in mean dispersal distance between center and edge increases linearly with patch radius, but the intensity of local adaptation depends on the dispersal kernel. Except for global and nearest neighbour dispersal, the evolved spatial pattern are qualitatively similar for both, mean dispersal probability and distance. We conclude, that inspite of the gene-flow originating from dispersal local adaptation of dispersal strategies is possible if a habitat is of sufficient size. This presumably holds for any realistic type of dispersal kernel.     

A communication-based spatial model of antipredator vigilance

Many animals spend their lives performing two often mutually exclusive tasks: feeding and watching out for predators (anti-predator vigilance). There have been many theoretical and empirical studies investigating this trade-off, especially for birds. An important characteristic of a flock of real birds is the area occupied by the flock. Individuals feeding close together experience increased competition so that the feeding rate decreases. Widely spaced individuals may suffer a loss in vigilance efficiency, since communication between individuals is more difficult, such that the predation risk increases. A vigilance model is developed which allows birds to control their spacing (and so the area of the flock) as well as their vigilance rate. The best strategy for the birds is found under a variety of environmental conditions, under the assumption that each individual acts selfishly to maximize its own fitness.     

相关风险因子对高原鼠兔摄食行为的影响

研究了捕食风险环境中集和洞口距离对高原鼠兔摄食行为的影响。结果表明,集群数量的增加不仅降低了警觉行为,同时也减少了摄食行为,在高风险环境中,集群为1时的取食行为强度最大,低风险环境中,为0时最大,警觉行为主要出现在距洞口2m的范围内,其行为强度与洞口踪影职责负相关,当洞口距离大于3m时,风险处理区的高原鼠兔几乎无警觉行为出现,且该处理区的取食区域几乎压缩的洞口旁,研究结果表明,在捕食风险环境中,高原鼠兔摄食行为与集群和洞口距离之间具有复杂的关系,其行为决策反映了降低风险与摄取食物间的权衡,行为目标是在降低捕食风险的同时尽可能地取食食物。