季节、性别和群体大小对鹅喉羚警戒行为的影响

动物通过集群降低个体警戒时间,从而增加采食等行为时间,这种现象被称为" 群体效应" 。除群体大小
外,社会及环境因子如季节与性别也可能影响个体警戒水平。本文于2007 年至2009 年在新疆卡拉麦里山有蹄类
野生动物保护区采用焦点动物取样法,通过测定鹅喉羚警戒行为比例、平均每次警戒持续时间及10 min内警戒频
率,研究了季节、性别及群体大小对鹅喉羚个体警戒水平的影响,并验证群体效应。结果表明:由于季节更替
而产生的生物量的变化对鹅喉羚个体警戒水平无显著影响;在秋冬季除雄性外,鹅喉羚个体警戒水平均有随群
体增大而降低的趋势,但这种趋势并不十分明显。性别则显著影响其警戒水平,夏季由于雌性羚羊处于哺乳期,
因而其警戒水平显著高于雄性(P < 0.05);受繁殖行为影响,冬季雄性比雌性警觉性更高(P > 0. 05);春秋季
两性间警戒水平没有差异(P >0.05)。不同生理周期是导致鹅喉羚育幼期雌性及繁殖期雄性个体警戒水平发生
显著变化的重要因子。     

动物的警戒行为——回顾及展望

警戒行为是动物对群体内部或外部刺激的行为反应,主要表现为抬头环视周围环境,具有反捕食和社会性监视功能,有利于个体适合度的增加。诸多研究表明,多种因素可以影响到动物警戒行为的强度和模式。内部因素主要包括个体的年龄、性别、所处空间位置和群体大小等,外部因素主要是捕食风险和人类活动的干扰。近年来,动物警戒同步性研究引起了广泛关注。同步警戒和协调警戒对立于独立警戒而存在,强调个体能够受到周围个体警戒状态的影响,继而表现出相同或相反的行为,并且其强度与群体大小等因素有关。在今后的研究中,可以在警戒行为影响因素和同步性方面进行更加深入而全面的探索。此外,警戒行为的个体差异和时空变化等也将是重要的研究方向。     

新疆科克森山盘羊季节性昼间行为时间分配和活动节律

2012年8月至2014年8月,采用目标取样法在新疆准噶尔界山西北部的科克森山区域对盘羊的行为进行了调查。盘羊行为被划分为采食、警戒、移动、休息和"其他"5种类型。共监测有效个体1197只,累计获得有效行为样本2068个,采食和休息是主要行为,平均时间所占百分比为43.1%和25.5%。除了春秋季雌性盘羊,其余季节盘羊5种行为间行为时间分配差异显著或极显著(P0.05或P0.01);除了冬季雌雄盘羊休息行为的平均时间差异不显著外,其余各季节雌雄盘羊各种行为差异极显著(P0.01);季节因素对雄性和雌性盘羊5种行为的平均时间分配均存在极显著影响(P0.01)。不同季节盘羊行为的日活动节律基本呈采食-休息-采食的活动模式,受地域环境因素和人为活动的影响,其活动模式具有自身的特异性。盘羊的行为时间分配和活动节律受食物资源、温度、同类竞争、天敌、繁殖周期和人类活动等影响。与其他研究结果对比后发现,不同区域的盘羊表现出不同的行为特征,表明动物的行为特征是动物本身长期进化和适应周围环境综合作用的结果。     

条纹短攀鲈的雄性个体大小、雌性偏好以及雄性之间的竞争

本研究考察了条纹短攀鲈(Trichopsis vittata)雄性个体的身体大小对雌性个体性偏好的影响以及对雄性之间竞争的影响。本研究设立了两种处理来研究雌性个体的性偏好以及雄性之间的竞争。在一种处理中,放入同样大小的雄性个体,而在另一种处理中,放入大小不同的雄性个体。结果表明,雄性个体的大小不影响雌性条纹短攀鲈的性偏好。在对雄性之间竞争的研究中,发现个体较大的个体获胜的次数比较多。在个体大小相同的处理中,雄性之间打斗的持续时间比另一种处理中的打斗持续时间长。对这两种处理中不同对抗行为的比较表明,个体大小相同处理组中的发声、咬、以及总体对抗行为发生得更为频繁。对个体大小不同处理组中的大、小个体的进一步观察也揭示出体型大的个体比体型小的个体有更多的追逐行为和更少的逃脱行为。这些结果表明,在雄性个体比雌性个体大的物种中,体型的性两型可能与雄性之间的竞争有关,而不是与雌性偏好有关。     

麻雀幼鸟和成鸟逃逸距离的比较

逃逸是鸟类常用的反捕食手段。自然选择会优化鸟类的逃逸距离,以便在躲避被捕食风险和保持能量之间做出权衡。理论模型预测动物个体可依据期望寿命来调整自身的行为：期望寿命短的个体倾向于风险偏好,即逃逸距离短;而期望寿命长的个体倾向于风险回避,即逃逸距离长。同时,逃逸距离还受到外部因素（如人为干扰强度、鸟类群体大小）的影响。本研究在北京城区收集了麻雀（Passer montanus）145只成鸟和75只幼鸟的逃逸距离数据,发现麻雀成鸟比幼鸟有着更长的逃逸距离,且成鸟更可能是群体中首先逃逸的个体。此外,研究发现麻雀的逃逸距离随着人为干扰强度的增加而降低,随着群体大小增加而增加。本研究结果符合理论预期：存活率高的类群（成鸟）有着更长的逃逸距离。     

群体规模对越冬灰鹤警戒行为的影响

2005年1—3月,在云南丽江拉市海就群体规模对越冬灰鹤(Grusgrus)警戒行为的影响进行了研究。用扫描取样记录群体的规模和警戒个体的数量、用焦点取样记录群体中个体警戒行为的频次和持续时间,结果显示灰鹤群体和个体的警戒力均随群体规模增加而降低,但集群个体数超过30只后,群体警戒力便不会再下降(P>0.05)、成体的警戒持续时间也会增加(P<0.01);当群体规模超过50只后,成体的警戒频次也会上升(P<0.05)。推测亚成体维持低警戒的群体规模上限值要高于成体,单从警戒行为分析,20—30只个体的集群可能代表越冬灰鹤的最适群体大小。     

蒙古野驴的警戒防御行为

2004 年6 ～12 月和2005 年3 ～7 月,在内蒙古自治区包头市达尔罕茂明安联合旗中蒙边境地区和巴彦淖尔市乌拉特梭梭林—蒙古野驴国家级自然保护区,采用瞬时扫描取样法,观察了蒙古野驴的警戒防御行为,共观察到蒙古野驴132 群次。蒙古野驴的防御行为可分为警觉和发现、预警、矛盾(犹豫)、示威、逃跑、护幼、诱发和争斗8 种方式,观察记录到16 种体姿。用独立样本t-test 检验,对成体蒙古野驴雌雄个体昼间预警行为时间分配进行差异显著性检验,结果发现雄性昼间用于预警的时间明显高于雌性。群体中所有个体均能通过体姿、动作和声音发出示警信号,逃跑是蒙古野驴躲避敌害的主要方式。野外观察记录到94 次警戒防御行为。统计结果表明,蒙古野驴的安全距离与调查者所使用交通工具有关,并与交通工具的运动速度呈正相关,因此提出了一个描述蒙古野驴安全距离与威胁源运动速度之间关系的模型。蒙古野驴具有激烈的种内争斗和攻击行为,激烈的争斗甚至导致被攻击个体死亡。      

四川察青松多白唇鹿国家级自然保护区白唇鹿繁殖期警戒行为

白唇鹿Cervus albirostris是青藏高原及其周边地区特有种,国家Ⅰ级重点保护野生动物。为探讨白唇鹿对环境影响的警戒响应,于2016年9—10月采用焦点取样法对四川察青松多白唇鹿国家级自然保护区白唇鹿繁殖期的警戒行为进行了观察,以探究影响白唇鹿个体警戒水平的因素,并利用白唇鹿警戒间隔片段数据来验证Pulliam警戒模型中顺序随机性的先提假设,即任意2次连续扫视间隔片段的时间长短是随机的,与前一次警戒间隔的长短无关。结果表明:白唇鹿的年龄性别对个体警戒水平有显著影响,幼崽警戒水平高于成体,雌性高于雄性;人类干扰程度和个体所处空间位置对警戒水平的影响不显著;93.4%的警戒间隔片段均为随机分布,表明白唇鹿警戒行为模式符合Pulliam模型中关于警戒间隔顺序随机性的假设。     

笼养褐马鸡繁殖期与非繁殖期行为观察与比较

2014年4—10月,利用焦点个体取样法与连续记录法对北京市野生动物救护繁育中心饲养的褐马鸡Crossoptilon mantchuricum繁殖期以及非繁殖期的行为时间分配和活动节律进行了研究。统计了不同性别与不同季节的行为时间分配差异。在繁殖期,雌性褐马鸡的取食时间大于雄性,而雄性褐马鸡的移动、警戒行为时间大于雌性。在非繁殖期,雌雄褐马鸡行为差异变小,但雄性褐马鸡的警戒行为仍然较多。     

群体规模对越冬灰鹤警戒行为的影响

2005年1—3月，在云南丽江拉市海就群体规模对越冬灰鹤(Grus grus)警戒行为的影响进行了研究。用扫描取样记录群体的规模和警戒个体的数量、用焦点取样记录群体中个体警戒行为的频次和持续时间，结果显示：灰鹤群体和个体的警戒力均随群体规模增加而降低，但集群个体数超过30只后，群体警戒力便不会再下降(P>0.05)、成体的警戒持续时间也会增加(P<0.01)；当群体规模超过50只后，成体的警戒频次也会上升(P<0.05)。推测亚成体维持低警戒的群体规模上限值要高于成体，单从警戒行为分析，20—30只个体的集群可能代表越冬灰鹤的最适群体大小。     

Vigilance and group size in impala (Aepyceros melampus Lichtenstein): a study in Nairobi National Park, Kenya

One of the advantages of living in groups is that individuals may need to be less vigilant, allowing them more time for other important activities, such as foraging. This relationship between group size and per cent time spent being vigilant was investigated by observing impala in Nairobi National Park, Kenya. Three types of individual were observed: females, territorial males and bachelor males. Only females showed the predicted negative relationship between per cent vigilance and herd size. Both types of male showed no significant change of vigilance with increasing group size. There was no difference in levels of vigilance in open or closed habitats and no difference in vigilance between herds ‘alone’ and herds with other species that might have provided ‘extra eyes’.     

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.     

Theory and method in studies of vigilance and aggregation

Predation is considered one of the most important selective pressures on free-ranging animals. Our understanding of it derives mainly from studies of individual vigilance (visual scanning of the surroundings beyond the immediate vicinity) and aggregation in prey. Vigilance bears a direct relationship to aggregation, because animals in groups may rely on associates for early warning of danger. This review addresses the relationship between vigilance and aggregation with particular attention to the prediction that individual vigilance declines with increasing group size. Contrary to most other animals studied, primates do not support the prediction. Exploring this, I examined the assumptions underlying vigilance theory in the light of primate behaviour. First I tested whether manual harvesting and upright processing of food as seen among primates might permit them to feed and scan simultaneously. I found no support for this idea. Next I examined the targets of primate vigilance and found that one component (within-group vigilance) might explain the differences between primates and other animals. Finally, I evaluated whether individual primates in large groups face a lower risk of predation than those in small groups. A conclusion was impossible, but by separating group-level from individual-level risk, I was able to identify several common circumstances in which group size would not predict individual risk or vigilance. These circumstances arose for primates and nonprimates alike. I concluded that the relationship of vigilance to aggregation is not straightforward. The absence of a group-size effect on vigilance among primates is probably due to functional differences in vigilance behaviour or safety in groups, not to methodological differences. Furthermore, future work on animal vigilance and aggregation must fully consider both the targets of glances, and the assumption that larger groups are safer from predators. I predict that animals will not relax vigilance in larger groups if conspecific threat increases with group size. Group size will not predict individual risk of predation nor individual vigilance rates when predators do not rely on surprise, or when predators select a small subset of highly vulnerable group members. Copyright 2000 The Association for the Study of Animal Behaviour.     

Vigilance and Collective Detection of Predators in Degus (Octodon degus)

Individuals of social and partially social species typically reduce their vigilance activity when foraging in groups. As a result, per capita risk of predation decreases and individuals allocate more time to foraging and other fitness rewarding activities. Reduction of per capita risk is hypothesized to occur because there are more individuals to detect potential predators. If so, collective (i.e. total) vigilance is expected to increase with foraging group size. Increased surveillance during group foraging may occur if group members scan independently of one another, or sequentially to avoid the overlapping of their vigilance bouts. Intriguingly, such coordinated vigilance assumes that individuals monitor not only the presence, but the vigilance behaviour of group mates. We used seasonal records on time budget and grouping patterns of individually marked degus (Octodon degus), a social rodent, to examine if (a) individual vigilance decreases and/or foraging increases with group size, (b) collective vigilance increases with group size and (c) foraging degus coordinate their vigilance. When foraging, degus decreased their individual vigilance and increased their foraging time when in larger groups. Despite this, degus in larger groups increased their collective vigilance, supporting the hypothesis that socially foraging degus decrease predation risk through an improved ability to detect and escape potential predators. Additionally, patterns of collective vigilance suggested that degus scan independently of each other and so, they do not coordinate their vigilance to prevent its temporal overlapping. This finding does not support that foraging degus monitor the vigilance activity of group mates.     

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.     

Human Activity Dampens the Benefits of Group Size on Vigilance in Khulan (Equus hemionus) in Western China

Animals receive anti-predator benefits from social behavior. As part of a group, individuals spend less time being vigilant, and vigilance decreases with increasing group size. This phenomenon, called “the many-eyes effect”, together with the “encounter dilution effect”, is considered among the most important factors determining individual vigilance behavior. However, in addition to group size, other social and environmental factors also influence the degree of vigilance, including disturbance from human activities. In our study, we examined vigilance behavior of Khulans (Equus hemionus) in the Xinjiang Province in western China to test whether and how human disturbance and group size affect vigilance. According to our results, Khulan showed a negative correlation between group size and the percentage time spent vigilant, although this negative correlation depended on the groups’ disturbance level. Khulan in the more disturbed area had a dampened benefit from increases in group size, compared to those in the undisturbed core areas. Provision of continuous areas of high-quality habitat for Khulans will allow them to form larger undisturbed aggregations and to gain foraging benefits through reduced individual vigilance, as well as anti-predator benefits through increased probability of predator detection.     

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.     

The effects of group size and reproductive status on vigilance in captive Callithrix jacchus

Previous work on mammals and birds has often demonstrated a negative relationship between group size and individual vigilance. However, this relationship has received only weak support in nonhuman primates. This result may be due to the failure to distinguish different forms of vigilance such as antipredatory vigilance and social monitoring. Here, we tested the effects of group size, reproductive status (breeding vs. nonbreeding), and sex on antipredatory vigilance and social monitoring in captive common marmosets (Callithrix jacchus). Behavioral observations using one-zero sampling were conducted on adult members of three captive groups of small, medium, and large size. Data were analyzed using a series of general linear models (GLMs) analyses of covariance (ANCOVAs). We found an overall negative group size effect on antipredatory vigilance and that breeders, especially breeding males, were significantly more vigilant than nonbreeders. Conversely, we found that social monitoring increased with group size. Unlike the results for antipredatory vigilance, neither breeders and nonbreeders nor males and females differed in their amounts of social monitoring. However, the effect of group size appeared to differ for nonbreeding males compared to all other adults. Our results generally support the idea that individuals in larger groups are safer with breeding males likely playing a prominent role in protection from predation. The increase in social monitoring may be related to increased reproductive competition with the presence of adult offspring, but future studies need to clarify the target of social monitoring in both breeders and nonbreeders. Overall, the study underlines the importance of distinguishing different forms of vigilance and other factors as they may confound the effects of group size on antipredatory vigilance.     

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。     

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.