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.     

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.     

Sexual,Seasonal and Group Size Differences in the Allocation of Time between Vigilance and Feeding in the Greater Rhea,Rhea americana

We studied the effect of sex and group size on the proportion of time a greater rhea, Rhea americana, allocates to vigilance and feeding during the breeding and the non-breeding seasons. We analysed 175 records of focal animals that were feeding alone or in groups of 2 to 26 birds. In both seasons, males spent more time in vigilance and less time in feeding than females. Both sexes spent more time in vigilance and less time in feeding during the breeding season. Sexual and seasonal differences in vigilance were the result of different mechanisms. Males had shorter feeding bouts than females but there were no sexual differences in the length of the vigilance bouts. On the contrary, seasonal differences were the result of males and females having longer vigilance bouts during the breeding season but there were no seasonal differences in the length of the feeding bouts. During the non-breeding season, individual vigilance was higher in rheas foraging alone than in groups. In this case, solitary birds had longer vigilance and shorter feeding bouts than birds foraging in groups. We discuss the possible effect of intragroup competition and food availability on the allocation of time between feeding and vigilance in this species.     

The effect of group size on vigilance in Ruddy Turnstones Arenaria interpres varies with foraging habitat

Foraging birds can manage time spent vigilant for predators by forming groups of various sizes. However, group size alone will not always reliably determine the optimal level of vigilance. For example, variation in predation risk or food quality between patches may also be influential. In a field setting, we assessed how simultaneous variation in predation risk and intake rate affects the relationship between vigilance and group size in foraging Ruddy Turnstones Arenaria interpres. We compared vigilance, measured as the number of ‘head‐ups’ per unit time, in habitat types that differed greatly in prey energy content and proximity to cover from which predators could launch surprise attacks. Habitats closer to predator cover provided foragers with much higher potential net energy intake rates than habitats further from cover. Foragers formed larger and denser flocks on habitats closer to cover. Individual vigilance of foragers in all habitats declined with increasing flock size and increased with flock density. However, vigilance by foragers on habitats closer to cover was always higher for a given flock size than vigilance by foragers on habitats further from cover, and habitat remained an important predictor of vigilance in models including a range of potential confounding variables. Our results suggest that foraging Ruddy Turnstones can simultaneously assess information on group size and the general likelihood of predator attack when determining their vigilance contribution.     

Collective Vigilance in the Wintering Hooded Crane: The Role of Flock Size and Anthropogenic Disturbances in a Human‐Dominated Landscape

Vigilance achieved at the group level, known as collective vigilance, can enhance the ability to assess threats and confer benefits to gregarious prey species. Examining the factors that influence collective vigilance and exploring how individual vigilance is organized at the group level can help to understand how prey groups respond to potential threats. We quantified collective vigilance and determined its temporal pattern in a natural wintering population of the hooded crane Grus monacha in the Shengjin Lake reserve in China. We examined the role of flock size and anthropogenic disturbances in the human‐dominated landscape on collective vigilance and level of synchronization. The proportion of time during which at least one individual scanned the surroundings (collective vigilance) increased with flock size and was higher in the more disturbed buffer zone of the lake. Synchronization of vigilance occurred more frequently in the smaller flocks but was not related to the risk of disturbance. Synchronization implies that individuals tend to monitor and copy the vigilance of neighbors. In the degraded wetlands, the wintering hooded crane benefits from foraging in groups and synchronizing their vigilance in response to human disturbances.     

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.     

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.     

Relation of group size and daily activity patterns to southern lapwing (Vanellus chilensis) behaviour

Behavioural patterns of birds commonly vary according to flock size and daily activity pattern. Southern lapwing behaviours and their relation with flock size were studied, as well as the relationship between the frequency of behaviours and the period of the day. Results showed that the proportion of time spent in foraging and vigilance was higher during the morning, when small groups were more common, and decreased from midday on, when group size increased. Maintenance and inactivity (sleeping) behaviours presented the opposite pattern. Correlation between flock size and period of the day, and their similar effects on bird behaviour may be evidence that groups of different sizes could have different functions throughout the day. Our results show that southern lapwings seem to form smaller flocks for feeding in the beginning of the day and larger flocks later for different activities (e.g., maintenance and sleeping). In this sense, it is possible that group size variation throughout the day is related to different demands for specific behaviours (functions) of groups, according to the daily activity patterns of the birds.     

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.     

Group‐Size Effect on Vigilance and Foraging in a Predator‐Free Population of Feral Goats (Capra hircus) on the Isle of Rum,NW Scotland

Many previous studies have found that as group size increases, individual vigilance levels decrease and forage intake increases (group‐size effect), but few such studies have considered the impact of within‐group interactions and other confounding factors on the direction of group‐size effects. A free‐ranging population of feral goats (Capra hircus), with little predation threat, was studied on the Isle of Rum (northwest Scotland), from Jun. to Nov. 2000, to investigate the effects of group size on individual vigilance levels and foraging efficiency after taking into account the effect of confounding factors (e.g. sex, season, time of day, habitat, predation risk) and within‐group interactions (via changes in movement rates while feeding). Our results show that, while group size exerted a negative influence on individual vigilance levels and a positive effect on movement rate, foraging efficiency never increased with group size and even decreased at certain times of day. There was no sex difference in individual vigilance in feral goats, but foraging efficiency was higher in females than in males. Goats were more vigilant in fall than in summer. The results imply that the benefits for foraging obtained from the reduced vigilance level in larger groups may be constrained or offset by increased interaction (or competition) within larger groups even in a population that faces negligible predation risk.     

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.     

反捕食不能充分解释獐的集群行为(英文)

我在江西鄱阳湖国家自然保护区研究了獐 (Hydropotesinermis)的集群行为并检验了集群是动物反捕食对策的假说。如果这个假说成立 ,我们则可以做三个预测 :(1)集群大小应与植物覆盖度成负相关 ;(2 )集群动物比单独活动的动物有更多的时间取食 ;(3)集群动物与单独活动的动物相比 ,用于警戒的时间较少。结果表明 ,獐的集群在交配季节最大而在产仔季节最小 ,獐在短草期比在高草期倾向于形成较大集群。因此 ,本研究支持了第一个预测。但是 ,集群的獐和单独活动的獐在取食和警戒时间分配上并无差异。因此 ,本研究不支持第二和第三个预测。时间收支 (timebudget)分析显示 ,在非交配季节 ,带幼仔的雌性用于取食的时间较少但走动频繁。在交配季节 ,与单独活动的雌性相比 ,与雄性在一起的雌性用于取食的时间较少 ,而单独活动的雄性却比与雌性在一起的雄性花较多的时间用于观望寻找雌性。鉴于獐集群行为的复杂性 ,本研究认为 ,獐并不只因反捕食而集群 ,其它社会和生态因子在决定獐的集群行为中也很重要。     

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.     

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.     

The effects of human presence, flock size and prey density on shorebird foraging rates

Animals may alter their foraging behaviour in the presence of humans because they perceive humans as potential predators. In this study I determined whether people caused shorebirds to reduce feeding rates at a stopover site in coastal British Columbia, Canada. I controlled for prey density and flock size because these variables may influence both the foraging rates as well as the effect of human disturbance on feeding efficiency. Semipalmated plovers decreased feeding rates when there were more people on the beach (multiple regression: F_1,15=5.86, b=0.59, P=0.029, R²=37.6%). For least sandpipers, the effect of human densities on feeding rates depended on flock size (F_1,21=5.97, P=0.023) and amphipod availability (F_1,21=4.98, P=0.037). This study demonstrated the importance of measuring subtle behavioural changes in foraging rates along with key ecological variables in order to assess the true impact of human disturbance on migratory shorebirds.     

Vigilance behavior during the birth and lactation season in naturally occurring ring-tailed lemurs (Lemur catta) at the Beza-Mahafaly Reserve,Madagascar

I examined the vigilance behavior of adult males and females in two groups of ring-tailed lemurs(Lemur catta)during the birth and lactation season at the Beza-Mahafaly Reserve, southwestern Madagascar. I found no sex difference with respect to the rates of overall vigilance, rates of vigilance toward a potential predator or unfamiliar sound, or rates of vigilance toward conspecifics from other social groups, nor were there sex differences in the percentage of time spent vigilant in any of the above categories. Higher-ranking females were vigilant significantly more often toward predators or potential predators than lower-ranking females were. I detected no relationship between vigilance behavior and dominance rank among adult males. The alpha female in each group exhibited significantly more vigilance behavior than all other members of her group. It was predicted that males should exhibit more vigilance behavior than females do, particularly during the birth and lactation season, when predator pressure is high, if they are benefiting females in this respect. I discuss the results in the context of this prediction and in terms of how ring-tailed lemur males benefit females, and why they may be tolerated in social groups.     

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.     

暂时与群体分离的个体藏马鸡的反捕食警戒

通过与同伴分担捕食风险 ,生活在稳定群体中的个体动物可以获得长期的适合度利益 ,但同时它们不得不承担食物竞争所带来的潜在代价。这种代价常常取决于食物资源的类型。当好的食物资源出现时 ,一些个体可以离开群体而独享这种资源。了解这些临时游离者如何组织其反捕食行为 ,在进化生态学上是有意义的。藏马鸡 (Crossoptilonharmani)是西藏雅鲁藏布江中游高山灌丛植被的一种典型的非繁殖季节集群鸟类。野外观察表明 ,为了独享好的食物资源 ,一些个体常常远离当前群体的活动范围。分离事件更可能发生于大的群体 ,但其发生率与参与者的数量呈负相关 ;而参与分离的个体愈多 ,分离持续的时间就愈长。分离者的个体警惕水平随着临时群体大小的增加而下降 ,遵从在其它自然大小鸟类群体所发现的一般性规律。分离行为的发生和持续时间被认为是个体对当前食物回报和捕食风险进行权衡的结果。这种利益 -代价权衡也可以解释藏马鸡所具有的强烈集群行为     

The advantages of social foraging in downy woodpeckers

I investigated the advantages gained by downy woodpeckers (Picoides pubescens) which join mixed-species winter flocks. Woodpeckers foraging alone showed high levels of vigilance as measured by head-cocking rates, and low feeding rates. Woodpeckers foraging with one or two flock members showed intermediate rates of head-cocking and feeding, while woodpeckers foraging with flocks of three or more birds showed low head-cocking rates and high feeding rates. Although local enhancement and copying may contribute to the woodpeckers' increased foraging efficiency in a flock, these do not appear to be the main factors. As downy woodpeckers spend less time on vigilance, they devote more time to foraging, thereby increasing their foraging efficiency     

年龄、集群、生境及天气对鄱阳湖白鹤越冬期日间行为模式的影响

于2012年1—3月和2012年12—2013年3月,采用瞬时扫描法研究了鄱阳湖国家级自然保护区越冬白鹤(Grusleucogeranus)的昼间时间分配模式和日活动节律,分析了年龄、集群大小、生境类型和天气对白鹤越冬行为的影响。结果表明,白鹤越冬期的主要行为有觅食、警戒、理羽、游走、休息和争斗。其中,觅食是鄱阳湖越冬白鹤主要行为,占其昼间活动时间的82.94%;警戒占11.94%,理羽占3.52%,游走占1.88%,休息占0.15%,争斗占0.02%。白鹤在越冬期间的觅食、游走、休息和争斗行为存在明显的昼间节律性,其中游走、觅食和争斗行为在每天的上午和下午各存在1个高峰,休息行为在中午出现一个高峰;警戒和理羽行为则没有显著的昼间节律变化。年龄、集群大小、天气和生境对白鹤的越冬行为具有显著的影响:幼鹤取食的时间显著多于成鹤,而警戒时间则显著的小于成鹤;与家庭群中的个体相比,集体群中的个体取食和争斗时间显著增加,警戒时间显著减少;在天气晴朗时,白鹤的警戒时间显著大于阴天的警戒时间,取食时间则显著少于阴天时白鹤的取食时间;白鹤在大风天气显著增加理羽行为;在有雾的天气会显著增加取食时间,减少警戒时间。在草洲和浅水两种生境中,白鹤的游走和理羽行为存在显著差异,而觅食和警戒行为没有显著的差异,这可能源于草洲和浅水生境相互毗邻,并且具有相同程度的人为干扰。GLM分析结果表明,年龄和集群大小、年龄和生境、集群大小和天气、集群大小和生境、天气和生境对白鹤昼间时间分配模式的影响存在显著交互效应。