Influences of sex,group size,and spatial position on vigilance behavior of Przewalski’s gazelles

Group-living animals may need to spend less time being vigilant, consequently, having more time for other important activities such as foraging (i.e., group size effect). Przewalski’s gazelle (Procapra przewalskii) is a group-living social animal, and a study was conducted in Qinghai Province of China during June–August 2006 by using a continuous focal sampling method to investigate the influences of group size, sex, within-group spatial position, and nearest-neighbor distance on individual vigilance level (defined as scanning frequency per minute). Male gazelles were more vigilant than females. The gazelle’s vigilance level decreased with group size (group size effect), but only for females. The individuals at the central positions within a group were less vigilant than those at the peripheral positions, but the nearest-neighbor distance did not have any significant influence on the individual vigilance level. Our results support the hypotheses of group size effect and edge effects, but the sexual difference in vigilance level and in the response to group size effect on vigilance suggests that there may be sexual difference in the function and targets of vigilance behavior of Przewalski’s gazelles, which warrants more investigation, with incorporation of within-group spatial position, to better understand the mechanism underlying the group size effect and edge effect.     

Vigilance in Przewalski's gazelle: effects of sex, predation risk and group size

Vigilance in social animals is often aimed at detecting predators. Many social and environmental factors influence vigilance, including sex, predation risk and group size. During the summer of 2007, we studied Przewalski's gazelle Procapra przewalskii , an endemic ungulate to the Qinghai-Tibet Plateau, to test whether and how these three factors affect vigilance. We distinguished groups consisting of males, mothers with lambs and females without lambs making observations on groups in the presence or absence of nearby predators. We assessed the group-size effect on vigilance and how this varied with levels of predation risk and sex. Males and mothers scanned longer and with a higher frequency than females without lambs. Individuals were more vigilant under direct predation threat. Although vigilance generally decreased with group size, the extent of the decrease was independent of predation risk and was not significant in males. The results suggest that mothers are more vigilant suggesting greater vulnerability and that males may have increased their vigilance to compete for higher social ranks. The positive correlation between vigilance and predation risk and the negative correlation between vigilance and group size are consistent with earlier findings, but we failed to find an interaction between group size and predation risk on vigilance perhaps because vigilance levels are low even in small groups, thus making similar vigilant upward adjustments in both small and large groups.     

Unpredictability of vigilance in two sympatric Tibetan ungulates

Vigilance is important for anti-predation, and different animals adopt different vigilance strategies. Instantaneous and sequential randomness in vigilance behavior are two main principles for the classic Pulliam model (1973). Given this context, we studied the vigilance behaviors in two wild cloven-hoofed animals, the Tibetan antelope (Pantholops hodgsonii) and the Tibetan gazelle (Procapra picticaudata) on Qinghai–Tibet Plateau, to explore if the two randomness principles work across species. The results showed that the distribution of inter-scan intervals of both Tibetan antelope and Tibetan gazelle followed the negative exponential distribution; inter-scans of both Tibetan antelope and Tibetan gazelle were unrelated with their previous scan, and most sequences of inter-scan intervals could be considered as random organized or unpredictable. In conclusion, the vigilance patterns of Tibetan antelope and Tibetan gazelle followed instantaneous randomness and sequential randomness of Pulliam model. A random vigilance strategy might be the best choice for Tibetan ungulates, and how to distinguish the social vigilance from anti-predator vigilance is an important issue for future research.     

The group size effect and synchronization of vigilance in the Tibetan wild ass

Vigilance behavior is considered as an effective strategy for prey species to detect predators.An individual benefits from living in a group by reducing the time spent being vigilant without affecting the probability of detecting a predator.However,the mechanism producing a decrease in vigilance with increasing group size is unclear.Many models of vigilance assume that group members scan independently of one another.Yet in recent studies,the other 2 patterns of vigilance,coordination and synchronization,were reported in some species.In 2 summers(2018 and 2019),we studied the group-size effect on vigilance and foraging of Tibetan wild ass in Chang Tang Nature Reserve of Tibet.We also tested whether individuals scan the environment independently,tend to coordinate their scans,or tend to synchronize their vigilance.The results showed that individuals decreased the time spent on vigilance with increasing group size,while increased the time spent foraging.Group members scanned the environment at the same time more frequently and there was a positive correlation between group members'behaviors,indicating that Tibetan wild asses tend to synchronize their vigilance.     

三种珍稀有蹄类动物的警戒行为数据

警戒行为是野生动物重要的反捕食策略之一, 警戒行为的研究已经成为行为生态学中继动物的采食行为、繁殖行为之外的另一热点研究领域。然而, 尽管研究人员至今已经在数百种鸟类、兽类中开展了各种类型的警戒行为学研究, 但相关研究的原始数据却鲜有共享。本文公开了利用焦点取样法采集的我国3种珍稀有蹄类动物警戒行为数据样本716个, 其中普氏原羚(Procapra przewalskii) 175个、藏原羚(P. picticaudata) 180个、麋鹿(Elaphurus davidiamus) 361个, 并同时公布了采集该行为学数据时的相关变量, 包括性别、群体类型、群体大小、捕食风险、人类干扰等。结果显示在上述3种有蹄类动物中, 均存在明显的集群效应, 且此效应与性别及捕食风险存在交互作用。本文尝试建立标准的行为学数据规范, 以期为未来的行为学研究数据的共享及深度挖掘提供可能。     

Interference competition and group size effect in sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) at salt licks

Competition has long been considered as a confounding factor of group size effect but the understanding of interference competition is rudimentary for the difficulty in disentangling interference competition from scramble competition adequately. Here, we analysed remote-camera video records of wild sika deer (Cervus nippon) at salt licks in southern China from March 1, 2006 to November 30, 2008 to investigate how interference competition and predation risk interacted on vigilance behaviour. Scramble competition is negligible at salt licks; therefore, we could focus our interest in interference competition. We used linear mixed model to compare vigilance, licking and aggression behaviours among females with and without fawn as well as males with different group sizes to identify the primary role of vigilance behaviour in sika deer. In total, 168 individuals were recorded and observation time was 2,733.04 min. We found that deer spent much time on vigilance and scanned frequently in spring and winter, and females with fawn spent more time on vigilance than females without fawn, suggesting vigilance for predation risk. Aggression ratio increased first and then decreased, while scan frequency continued to decline and then slightly increased when group size increased from two to seven, implying vigilance for interference competition. Our results suggested vigilance in sika deer was influenced by both predation risk and interference competition, but was mainly driven by predation risk even at sites with intense interference competition. Our results of interference competition shed some light on finding the underlying mechanism of group size effect in wild populations.     

Effects of reproductive status, social rank, sex and group size on vigilance patterns in Przewalski's gazelle

Background

Quantifying vigilance and exploring the underlying mechanisms has been the subject of numerous studies. Less attention has focused on the complex interplay between contributing factors such as reproductive status, social rank, sex and group size. Reproductive status and social rank are of particular interest due to their association with mating behavior. Mating activities in rutting season may interfere with typical patterns of vigilance and possibly interact with social rank. In addition, balancing the tradeoff between vigilance and life maintenance may represent a challenge for gregarious ungulate species rutting under harsh winter conditions. We studied vigilance patterns in the endangered Przewalski''s gazelle (Procapra przewalskii) during both the rutting and non-rutting seasons to examine these issues.

Methodology/Principal Findings

Field observations were carried out with focal sampling during rutting and non-rutting season in 2008–2009. Results indicated a complex interplay between reproductive status, social rank, sex and group size in determining vigilance in this species. Vigilance decreased with group size in female but not in male gazelles. Males scanned more frequently and thus spent more time vigilant than females. Compared to non-rutting season, gazelles increased time spent scanning at the expense of bedding in rutting season. During the rutting season, territorial males spent a large proportion of time on rutting activities and were less vigilant than non-territorial males. Although territorial males may share collective risk detection with harem females, we suggest that they are probably more vulnerable to predation because they seemed reluctant to leave rut stands under threats.

Conclusions/Significance

Vigilance behavior in Przewalski''s gazelle was significantly affected by reproductive status, social rank, sex, group size and their complex interactions. These findings shed light on the mechanisms underlying vigilance patterns and the tradeoff between vigilance and other crucial activities.     

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     

Sexual segregation in Tibetan gazelle: a test of the activity budget hypothesis

Tibetan gazelle Procapra picticaudata , a threatened endemic species on the Qinghai-Tibet Plateau, is a sexually dimorphic ungulate. Males are larger than females and both sexes live in different groups, except during the rutting season. In order to test the activity budget hypothesis, which was proposed to explain social segregation in ungulates, we studied the activity budget of male and female Tibetan gazelles during the summers of 2005 and 2006. The activity budget hypothesis predicts that females spend more time feeding and group members synchronize their activities more in the same-sex groups than in the mixed-sex groups. We found that females and males of Tibetan gazelle had different activity budgets; females spent significantly longer time feeding and comparably less time on other activities. Activity synchronization indexes between female groups and male groups were similar (female groups: 0.81±0.17, male groups: 0.80±0.19), whereas both of them were significantly higher than that of mixed-sex groups (0.58±0.29). These results suggest that although sexual segregation might be caused by multiple mechanisms, the activity budget hypothesis about sexual segregation is supported by our studies of Tibetan gazelle on the Qinghai-Tibet Plateau.     

Investigating Differences in Vigilance Tactic Use within and between the Sexes in Eastern Grey Kangaroos

Aggregation is thought to enhance an animal’s security through effective predator detection and the dilution of risk. A decline in individual vigilance as group size increases is commonly reported in the literature and called the group size effect. However, to date, most of the research has only been directed toward examining whether this effect occurs at the population level. Few studies have explored the specific contributions of predator detection and risk dilution and the basis of individual differences in the use of vigilance tactics. We tested whether male and female (non-reproductive or with young) eastern grey kangaroos (Macropus giganteus) adopted different vigilance tactics when in mixed-sex groups and varied in their reliance on predator detection and/or risk dilution as group size changed. This species exhibits pronounced sexual dimorphism with females being much smaller than males, making them differentially vulnerable toward predators. We combined field observations with vigilance models describing the effects of detection and dilution on scanning rates as group size increased. We found that females with and without juveniles relied on predator detection and risk dilution, but the latter adjusted their vigilance to the proportion of females with juveniles within their group. Two models appeared to equally support the data for males suggesting that males, similarly to females, relied on predator detection and risk dilution but may also have adjusted their vigilance according to the proportion of mothers within their group. Differential vulnerability may cause sex differences in vigilance tactic use in this species. The presence of males within a group that do not, or only partially, contribute to predator detection and are less at risk may cause additional security costs to females. Our results call for reexamination of the classical view of the safety advantages of grouping to provide a more detailed functional interpretation of gregariousness.     

青藏公路对藏羚羊、藏原羚和藏野驴活动的影响

2003年8月和2004年8月,在可可西里国家自然保护区通过实地调查,分析了青藏公路运营对藏羚羊(Pantholops hodgsoni)、藏原羚(Procapra picticaudata)和藏野驴(Equus kiang)行为活动的影响.结果表明:藏野驴对青藏公路形成了回避,其在距路基1 001～2 000和2 001～3 000 m区域内的种群密度显著高于0～500 m的区域(P＜0.05);青藏公路对藏羚羊、藏原羚的行为活动产生了一定程度的干扰,尤其是对藏羚羊,其在距路基0～500 m区域内的行为活动与距路基2000 m之外的区域具有极显著差异(P＜0.01).同时,因其群体数量大,个体通过公路所花费的时间长,需要很长的车辆行驶间隔才能通过公路,故受车辆运输的干扰最大,无法顺利通过公路.研究还发现,在白天的不同时间段,藏羚羊、藏原羚在公路附近的数量分布与各时间段内的车流量呈极显著负相关,这有利于其穿越公路,说明野生动物通过自身的适应和行为调节可以减少环境改变所造成的影响.     

青藏高原异地半圈养藏羚警戒行为的适应性研究

行为是反映动物应对环境变化的最直接形式。动物可以根据周围环境条件的变化以及自身的生理状况来调整行为,异地放养是保护珍稀动物的有效方法,但必然会对其行为产生影响。为了探讨藏羚(Pantholops hodgsonii)对异地环境的行为学适应,对异地圈养藏羚的警戒行为进行了不同季节间的比较研究,采用全事件记录法和焦点动物取样法,记录和统计了异地圈养藏羚在冷季和暖季的警戒行为,进而推测其对人类干扰的行为适应性。研究结果表明,雌性和雄性藏羚的警戒时间及警戒比例(警戒时间占全天活动时间的比例)在暖季存在显著差异(警戒时间:Z=4.36,P<0.05;警戒比例:Z=4.559,P<0.05),而在冷季则无差异(警戒时间:Z=0.001,P>0.05;警戒比例:Z=0.0014,P>0.05);而季节差异对雌、雄性藏羚的警戒时间、警戒比例均具有极显著的影响(雄性-警戒时间:F=31.758,P<0.01;警戒比例:F=21.768,P<0.01;雌性-警戒时间:F=14.98,P<0.01;警戒比例:F=11.05,P<0.01);但是季节和性别对藏羚警戒行为的影响没有交互作用(Z=?0.576,P>0.05)。这些结果提示异地圈养藏羚警戒行为的变化可能是对陌生环境适应的结果。     

Interactions among social monitoring,anti-predator vigilance and group size in eastern grey kangaroos

Group size is known to affect both the amount of time that prey animals spend in vigilance and the degree to which the vigilance of group members is synchronized. However, the variation in group-size effects reported in the literature is not yet understood. Prey animals exhibit vigilance both to protect themselves against predators and to monitor other group members, and both forms of vigilance presumably influence group-size effects on vigilance. However, our understanding of the patterns of individual investment underlying the time sharing between anti-predator and social vigilance is still limited. We studied patterns of variation in individual vigilance and the synchronization of vigilance with group size in a wild population of eastern grey kangaroos (Macropus giganteus) subject to predation, in particular focusing on peripheral females because we expected that they would exhibit both social and anti-predator vigilance. There was no global effect of group size on individual vigilance. The lack of group-size effect was the result of two compensating effects. The proportion of time individuals spent looking at other group members increased, whereas the proportion of time they spent scanning the environment decreased with group size; as a result, overall vigilance levels did not change with group size. Moreover, a degree of synchrony of vigilance occurred within groups and that degree increased with the proportion of vigilance time peripheral females spent in anti-predator vigilance. Our results highlight the crucial roles of both social and anti-predator components of vigilance in the understanding of the relationship between group size and vigilance, as well as in the synchronization of vigilance among group members.     

Influence of population density on group sizes in goitered gazelle (Gazella subgutturosa Guld., 1780)

We conducted our study in Ili depression, south-eastern Kazakhstan during 1981–1989 to investigate how group sizes and group class frequencies change with increasing population densities in goitered gazelles. In addition, we compared our study to data on group size and group class frequency of various goitered gazelle populations in Kazakhstan with very variable population densities. We found that mean group size was a more variable index than group class frequency. Population density had some effect on mean group sizes, but the strength of the influence was quite weak, and only in cases where densities of two populations varied more than sevenfold did group sizes start to change. Group class frequency was not correlated with population density at all. The impact of the yearly breeding cycle on group size was bigger than population density. The density-dependent response of goitered gazelle population was curvilinear in fashion, and it may be classified as intermediate between social-dwelling ungulate species, living in large groups and demonstrating continuous (linear) increases of group size with population density and those that are solitary or territorial ungulate species with no relationship between population size and group size, though the goitered gazelle population’s weak response was distinctively closer to the one of solitary ungulate species.     

Vigilance behavior of Pyrenean chamois Rupicapra pyrenaica pyrenaica: Effect of sex and position in the herd

The Pyrenean chamois Rupicapra pyrenaica pyrenaica is a mountain-dwelling ungulate with an extensive presence in open areas. Optimal group size results from the trade off between advantages (a reduction in the risk of predation) and disadvantages (competition between members of the herd) of group living. In addition, advantages and disadvantages of group living may vary depending on the position of each individual within the herd. Our objective was to study the effect of central vs. Peripheral position in the herd on feeding and vigilance behavior in male and female Pyrenean chamois and to ascertain if a group size effect existed. We used focal animal sampling and recorded social interactions when a focal animal was involved. With males, vigilance rate was higher in the central part of the group than at the periphery, probably due to a higher density of animals in the central part of the herd and a higher probability of being disturbed by conspecifics. With females, vigilance rate did not differ according to position in the herd. Females spent more time feeding than males, and males showed a higher frequency of the vigilance behavior than females. We did not observe a clear relationship between group size and vigilance behavior. The differences in vigilance behavior might be due to social interactions.     

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.     

Foraging strategy and predator avoidance behaviour: an intraspecific approach

Relationships between predator avoidance behaviour (scanning and flocking) and foraging were studied in Calidris alpina, to test predictions regarding the effect of foraging techniques on such behaviours. The scanning hypothesis predicts that individuals with a tactile hunting technique and individuals with a visual hunting technique (both continuous searchers) do not differ in any variable related to scanning behaviour. The flocking hypothesis predicts that visually hunting individuals witl tend to form smaller flocks than tactile-foraging individuals. The two continuous feeding strategies did not differ among individuals in vigilance rate, nor in vigilance time or mean scan duration. However, with respect to flocking behaviour, visual foragers differed from tactile foragers in foraging flock size. The relationships between flocking behaviour and foraging strategy are discussed. The pattern found at the intraspecific level are the same as those found at interspecific level.     

Group size effects on foraging and vigilance in migratory Tibetan antelope

Large group sizes have been hypothesized to decrease predation risk and increase food competition. We investigated group size effects on vigilance and foraging behaviour during the migratory period in female Tibetan antelope Pantholops hodgsoni, in the Kekexili Nature Reserve of Qinghai Province, China. During June to August, adult female antelope and yearling females gather in large migratory groups and cross the Qinghai-Tibet highway to calving grounds within the Nature Reserve and return to Qumalai county after calving. Large groups of antelope aggregate in the migratory corridor where they compete for limited food resources and attract the attention of mammalian and avian predators and scavengers. We restricted our sampling to groups of less than 30 antelopes and thus limit our inference accordingly. Focal-animal sampling was used to record the behaviour of the free-ranging antelope except for those with lambs. Tibetan antelope spent more time foraging in larger groups but frequency of foraging bouts was not affected by group size. Conversely, the time spent vigilant and frequency of vigilance bouts decreased with increased group size. We suggest that these results are best explained by competition for food and risk of predation.     

How group size affects vigilance dynamics and time allocation patterns: the key role of imitation and tempo

In the context of social foraging, predator detection has been the subject of numerous studies, which acknowledge the adaptive response of the individual to the trade-off between feeding and vigilance. Typically, animals gain energy by increasing their feeding time and decreasing their vigilance effort with increasing group size, without increasing their risk of predation ('group size effect'). Research on the biological utility of vigilance has prevailed over considerations of the mechanistic rules that link individual decisions to group behavior. With sheep as a model species, we identified how the behaviors of conspecifics affect the individual decisions to switch activity. We highlight a simple mechanism whereby the group size effect on collective vigilance dynamics is shaped by two key features: the magnitude of social amplification and intrinsic differences between foraging and scanning bout durations. Our results highlight a positive correlation between the duration of scanning and foraging bouts at the level of the group. This finding reveals the existence of groups with high and low rates of transition between activities, suggesting individual variations in the transition rate, or 'tempo'. We present a mathematical model based on behavioral rules derived from experiments. Our theoretical predictions show that the system is robust in respect to variations in the propensity to imitate scanning and foraging, yet flexible in respect to differences in the duration of activity bouts. The model shows how individual decisions contribute to collective behavior patterns and how the group, in turn, facilitates individual-level adaptive responses.     

The effects of mixed-species flocking on the vigilance of shorebirds: Who do they trust?

The benefit of reduction in individual anti-predatory vigilance with increasing group size has previously been demonstrated only within single species. Here the effect of mixed species flocking on vigilance is investigated in two species of wading bird wintering on rocky shores. Both turnstones, Arenaria interpres, and purple sandpipers, Calidris maritima, ‘share’ vigilance with conspecifics, but also with some other waders; the extent of sharing appears to depend on the relative size of, and habitat overlap with, the other species. Vigilance is not shared with much larger species, nor with those rarely found in the same areas. There is no sharing of vigilance between neighbouring birds unable to see each other. Variation in vigilance level results from an alteration in both rate of scanning and duration of single scans; these two variables are adjusted independently over the range of densities experienced. This is due to constraints on a high scanning rate (which may reduce feeding efficiency) and a short scan duration (which cannot be lower than the minimum required to take in the necessary information).