Vigilance and Activity Time-Budget Adjustments of Wintering Hooded Cranes,Grus monacha,in Human-Dominated Foraging Habitats

Due to loss and degradation of natural wetlands, waterbirds increasingly rely on surrounding human-dominated habitats to obtain food. Quantifying vigilance patterns, investigating the trade-off among various activities, and examining the underlying mechanisms will help us understand how waterbirds adapt to human-caused disturbances. During two successive winters (November-February of 2012–13 and 2013–14), we studied the hooded crane, Grus monacha, in the Shengjin Lake National Nature Reserve (NNR), China, to investigate how the species responds to human disturbances through vigilance and activity time-budget adjustments. Our results showed striking differences in the behavior of the cranes when foraging in the highly disturbed rice paddy fields found in the buffer zone compared with the degraded natural wetlands in the core area of the NNR. Time spent vigilant decreased with flock size and cranes spent more time vigilant in the human-dominated buffer zone. In the rice paddy fields, the birds were more vigilant but also fed more at the expense of locomotion and maintenance activities. Adult cranes spent more time vigilant and foraged less than juveniles. We recommend habitat recovery in natural wetlands and community co-management in the surrounding human-dominated landscape for conservation of the hooded crane and, generally, for the vast numbers of migratory waterbirds wintering in the middle and lower reaches of the Yangtze River floodplain.     

西藏拉萨越冬黑颈鹤家庭群的警戒同步性

以往关于动物警戒行为的研究多以经典随机独立模型为基础,然而,近期一些理论和实证研究则向该模型发出了挑战。根据警戒行为同步性程度的不同,集体警戒可能存在基于模仿的同步警戒和避免重复的协调警戒两种模式。研究了中国雅鲁藏布江自然保护区越冬黑颈鹤家庭群的警戒行为,通过独立样本t检验验证由于年龄和是否携幼所导致的个体警戒水平的差异,通过单因素方差分析比较不同类型家庭群的集体警戒水平,通过配对t检验比较集体警戒的实际观测值和独立假设下的预期值从而确定越冬黑颈鹤家庭群的警戒同步性。结果表明家庭群中的幼年黑颈鹤警戒行为时间显著低于成年黑颈鹤,但幼鹤的存在对成鹤的警戒水平没有显著影响。不同类型黑颈鹤家庭群的集体警戒水平没有差异,但并未如预期表现为同步警戒。警戒行为模式是动物在特定环境中的适应性表现,越冬黑颈鹤生境中不存在落后目标策略的捕食者和高强度的捕食风险,因而可能不足以诱发以降低捕食风险为主要功能的高强度的警戒行为同步性;同时,在资源匮乏的冬季对食物获取的优先性可能也削弱了对同步警戒的需求。     

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.     

Disturbance and predation risk influence vigilance synchrony of black‐necked cranes Grus nigricollis,but not as strongly as expected

Animals monitor surrounding dangers independently or cooperatively (synchronized and coordinated vigilance), with independent and synchronized scanning being prevalent. Coordinated vigilance, including unique sentinel behavior, is rare in nature, since it is time‐consuming and limited in terms of benefits. No evidence showed animals adopt alternative vigilance strategies during antipredation scanning yet. Considering the nonindependent nature of both synchronization and coordination, we assessed whether group members could keep alert synchronously or in a coordinated fashion under different circumstance. We studied how human behavior and species‐specific variables impacted individual and collective vigilance of globally threatened black‐necked cranes (Grus nigricollis) and explored behavior‐based wildlife management. We tested both predation risk (number of juveniles in group) and human disturbance (level and distance) effects on individual and collective antipredation vigilance of black‐necked crane families. Adults spent significantly more time (proportion and duration) on scanning than juveniles, and parents with juveniles behaved more vigilant. Both adults and juveniles increased time allocation and duration on vigilance with observer proximity. Deviation between observed and expected collective vigilance varied with disturbance and predation risk from zero, but not significantly so, indicating that an independent vigilance strategy was adopted by black‐necked crane couples. The birds showed synchronized vigilance in low disturbance areas, with fewer juveniles and far from observers; otherwise, they scanned in coordinated fashion. The collective vigilance, from synchronized to coordinated pattern, varied as a function of observer distance that helped us determine a safe distance of 403.75 m for the most vulnerable family groups with two juveniles. We argue that vigilance could constitute a prime indicator in behavior‐based species conservation, and we suggesting a safe distance of at least 400 m should be considered in future tourist management.     

Contagious fear: Escape behavior increases with flock size in European gregarious birds

Flight initiation distance (FID), the distance at which individuals take flight when approached by a potential (human) predator, is a tool for understanding predator–prey interactions. Among the factors affecting FID, tests of effects of group size (i.e., number of potential prey) on FID have yielded contrasting results. Group size or flock size could either affect FID negatively (i.e., the dilution effect caused by the presence of many individuals) or positively (i.e., increased vigilance due to more eyes scanning for predators). These effects may be associated with gregarious species, because such species should be better adapted to exploiting information from other individuals in the group than nongregarious species. Sociality may explain why earlier findings on group size versus FID have yielded different conclusions. Here, we analyzed how flock size affected bird FID in eight European countries. A phylogenetic generalized least square regression model was used to investigate changes in escape behavior of bird species in relation to number of individuals in the flock, starting distance, diet, latitude, and type of habitat. Flock size of different bird species influenced how species responded to perceived threats. We found that gregarious birds reacted to a potential predator earlier (longer FID) when aggregated in large flocks. These results support a higher vigilance arising from many eyes scanning in birds, suggesting that sociality may be a key factor in the evolution of antipredator behavior both in urban and rural areas. Finally, future studies comparing FID must pay explicit attention to the number of individuals in flocks of gregarious 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.     

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.     

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 responding to number of conspecifics among mixed groups of cranes in demilitarized zone

Numerous studies have addressed antipredatory benefits of mixed-species flocks of foragers, but studies on individual's vigilance as a function of group size are limited. In the Cheolwon area of the Korean Demilitarized Zone, vigilance of the subordinate White-naped cranes (Grus vipio) in 11 groups composed of conspecifics and the dominant Red-crowned cranes (Grus japonensis) was examined. Vigilance correlated negatively with group size due to negative correlation with the number of conspecifics, but not the dominant heterospecifics. This is consistent with the hypothesis that a decrease in vigilance in larger groups is due to antipredatory benefits from increased predator detection in larger groups (associated with the presence of a larger number of conspecifics). This suggested that the mechanism leads to canceling out of the otherwise expected antipredatory benefits to the subordinate species from the increased predator detection by larger group size (associated with larger number of dominants). This is one of only a few behavioral studies of these endangered crane species in the relatively inaccessible wintering area of international importance in the areas of high conservation value.     

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.     

Scanning pattern of greater rheas,<Emphasis Type="Italic"> Rhea americana</Emphasis>: collective vigilance would increase the probability of detecting a predator

Many models using vigilance to predict the probability of detecting an approaching predator assumes that prey scanning events should be produced at random. Consequently, the length of intervals among successive scans must follow a negative exponential distribution. We analyzed the scanning behavior of the greater rhea, Rhea americana, which is a gregarious, flightless bird, in eastern Argentina. We investigated whether individual and/or collective scanning departed from random and whether this departure varied with group size. We used two simulation models based on observed scanning sequences to assess the effectiveness of vigilance on the individual and collective level when faced with an opportunistic or stalking predator. The analysis of 59 behavioral sequences of wild greater rheas foraging solitary or in groups of two to six or more individuals revealed that the inter-scan length of individual sequences significantly departed from random. In contrast, inter-scan intervals for collective vigilance were shorter than individual ones, but only fit the random expectation for groups of two and five individuals. Models showed that collective vigilance could increase the probability of detecting a predator, thereby reducing their vulnerability, independent of whether the predator uses a stalking or opportunistic approaching strategy.     

云南省纳帕海自然保护区越冬黑颈鹤的集群特征

2004年10月-2005年5月,在云南纳帕海自然保护区采用定点扫描法对越冬黑颈鹤(Grus nigricollis)的集群类犁和集群大小进行了观察.结果表明黑颈鹤夜间集群夜栖,形成较大的夜栖群,平均群体大小为67.9只(16-157,n=17):按照有无灰鹤加入,又将其分为同种集群和混种集群两种类型,其中同种集群的黑颈鹤数量占整个越冬种群的65.3%.在白昼,黑颈鹤以家庭鹤、集群鹤及特殊群体3种类型活动,家庭鹤和集群鹤的平均大小分别为2.7只(2-4,n=145)和16.1只(3-65,n=1017).黑颈鹤的集群大小并不稳定,在日内和月份间均有明显变化(P=0.000＜0.05).存越冬期,最大集群形成于12月,其次为11月和1月;在日内,早上8时集群最大,随后减小并保持相对稳定,18时黑颈鹤开始向夜栖地靠拢,使得集群再次开始增大.随后观察中还发现,黑颈鹤的家庭解体过程开始于3月底,当幼鹤被成鹤驱逐离群后,逐渐加入集群鹤活动,从而使得家庭鹤和集群鹤的大小和组成发生改变.黑颈鹤的集群大小和组成受自身状况、种内关系、天气、食物等多种因素的共同影响,随时间和季节变动而发生变化,是对自身、种群和环境条件变化的综合反映.     

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。     

Vigilance Decreases with Time at Loafing Sites in Gulls (Larus spp.)

Understanding how animals partition effort between vigilance for predators and other conflicting activities has been a mainstay of animal behaviour research. Classical theories implicitly assume that animals alternate between high and low vigilance states over short timescales, but that average effort invested in vigilance is constant over an extended bout of such alternations. However, one recent model suggests that vigilance should be adjusted dynamically to short‐term changes in the perception of predation risk and would tend to decrease with time. Indeed, as time passes by without disturbances, perception of the need for vigilance should decrease and prey animals may allocate more time to competing activities. Here, we examined how the proportion of sleeping gulls (Larus spp.) in loafing groups changed over time. Sleeping gulls can only maintain low levels of vigilance against external threats (compared to alert individuals), and we predicted that the proportion of sleeping gulls at loafing sites should increase over time when no disturbances occur. Statistically significant changes in the proportion of sleeping gulls as a function of time occurred in the majority of sequences and an increase was observed significantly more often than predicted by chance alone. This temporal pattern cannot be caused by reduction in hunger levels because gulls are not feeding at loafing sites. The results indicate that vigilance can be adjusted dynamically in response to short‐term temporal changes in the perception of predation risk.     

Vigilance in Single- and Mixed-Species Groups of Tamarins (Saguinus labiatus and Saguinus fuscicollis)

By forming larger sizes of groups, individuals benefit from a decrease in vigilance, but the collective vigilance of the group as a whole is not compromised. We examined whether this group size effect is apparent in mixed-species groups of red-bellied tamarins (S. labiatus) and saddleback tamarins (S. fuscicollis) which form stable and permanent associations in the wild. We studied general vigilance and responses to hidden threatening stimuli in five captive groups of each species, while they were housed in single- and mixed-species groups. For vigilance, the individual rate was lower in the larger mixed-species groups than in the smaller single-species groups. In addition, the amount of time when at least one individual was vigilant was higher in mixed-species groups. This suggests that the tamarins alter their vigilance behavior in the presence of the other species. In response to hidden threats, both species performed brief vigilance checks and frequencies of checking did not differ in single- and mixed-species groups. However, both species had a significant reduction in the mean duration per check, and there was a reduced total amount of time spent vigilance checking in the mixed-species groups compared to the single-species groups, demonstrating the group size effect. Overall the mixed-species groups had a higher number and mean duration of checking than the smaller single-species groups. Given that the two species share a common set of predators, and respond to each other's alarm calls, these findings provide strong evidence that individuals of both tamarin species may be able to benefit from forming mixed-species groups via improved vigilance and monitoring of threats.     

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.     

Does vigilance always covary negatively with group size? Effects of foraging strategy

The possible effects of foraging strategy on the relationship between vigilance and group size were studied in three species of waders with different foraging strategies. I predict that (1) pause–travel species should show no relationship between scanning and group size, because these species scan for prey as well as for predators; (2) continuous-tactile foraging species should show a positive relationship between flock size and vigilance level, because in their large groups vigilance towards other birds could be used to avoid interference and aggression; and (3) continuous-visual searching species should show the general pattern of decreasing vigilance when group size increases as predicted by both the 'many eyes' and the 'predatory risk' hypotheses. Results support the predictions for the influence of foraging strategy on the relationship between vigilance level and group size. The mutual exclusion of foraging and scanning can determine the importance of the 'many eyes' hypothesis. Such exclusion seems to be determined by foraging strategy. The influence of foraging on scanning must be taken into account in future studies on the group-size effect. Electronic Publication     

苏北地区丹顶鹤越冬种群数量及栖息地分布动态变化

利用资料追踪和历史时期土地利用变化数据深入研究了有资料记载以来苏北地区丹顶鹤越冬种群数量变化与栖息地分布的时空响应。结果显示,20世纪80年代之初,丹顶鹤越冬种群广泛分布于苏北地区,尤其是长江中下游的内陆湖泊、沼泽及江苏沿海滩涂湿地;20世纪90年代之后丹顶鹤分布区域逐渐向沿海滩涂湿地转移,主要分布于盐城市的射阳县境内;进入21世纪,丹顶鹤分布区域集中于盐城国家级珍禽保护区核心区。与此同时,丹顶鹤越冬种群数量也经历了十分明显的变化过程,与20世纪90年代相比,数量减少50%以上。丹顶鹤越冬种群数量及栖息地分布变化与栖息地分布点周围的土地利用方式及人为干扰有关,栖息地面积丧失和景观破碎化是丹顶鹤越冬种群数量减少的主要原因。     

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.