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。     

散放麋鹿的警戒行为模式、警戒距离及其季节差异

警戒行为是动物对生存环境的风险或潜在风险的一种行为反应(Li ma and Zollner,1996;Treves,2000)。近年来关于警戒行为的研究正在成为热点,内容涉及了警戒行为模式、警戒行为与反捕食策略的关系和群体警戒等诸多方面(Li maand Zollner,1996;Beauchamp,2001;Hass andValenzuela     

散放麇鹿的警戒行为模式、警戒距离及其季节差异

警戒行为是动物对生存环境的风险或潜在风险的一种行为反应（Lima and Zollner,1996;Treves。2000）。近年来关于警戒行为的研究正在成为热点,内容涉及了警戒行为模式、警戒行为与反捕食策略的关系和群体警戒等诸多方面（Limaand Zollner,1996;Beauchamp,2001;Hassand Valenzuela,2002）,并形成了两个从不同侧面阐述警戒行为功能和影响因素的重要理论：“个体风险理论”和“群体大小效应理论”（Limaand Dill,1990;Quenette,1990;Roberts,1996;Burgerand Gochfeld,2000）。关于鹿类动物警戒行为的研究只见于狍（Capreolus pygargus,Rebyeta1．,1999）、白尾鹿（Odocoileus virginianus,Lingleand Wilson,2001）和黑尾鹿（Odocoileus hemionus,Lingleand Wilson,2001）。     

云南纳帕海越冬黑颈鹤日间行为模式与年龄和集群的关系

2006年10月—2007年5月,在云南省西北部纳帕海,采用路线调查结合瞬时扫描行为取样法,对越冬黑颈鹤(Grus nigricollis)种群的时间分配及其与年龄、集群和时间的关系进行了观察。结果说明,黑颈鹤越冬活动主要是以觅食为主,占日间时间的(76.81±9.1)％。越冬期间黑颈鹤日间行为的节律性较为明显,具有适应高寒气候的特点。集群形式对成鹤的行为有着显著影响,集群和家庭中活动的成鹤在觅食、警戒和争斗中存在显著差异(F1,76= 0.27、0.77, U= 279, P= 0.001—0.000)。年龄是影响鹤群行为的因素之一。幼鹤相比成鹤有较多的觅食时间和休息时间,警戒行为比例较低(F1,76= 0.04—2.59, U= 188—299, P= 0.006—0.000),且不受集群形式的影响。随着越冬期间的早、中、晚3个时期的环境变化,黑颈鹤的时间分配有显著变化(F2,36= 4.63—26.54, 22= 5.29—13.68, P= 0.0016—0.000)。不同越冬地的黑颈鹤行为存在差异。气候、食物资源和人为影响可能是造成这些差异的主要因素。     

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。     

鄱阳湖稻田生境中越冬灰鹤行为模式及其影响因素

2018年10月—2019年3月,采用焦点动物法研究了鄱阳湖区稻田生境中灰鹤(Grus grus)越冬期的行为模式及其影响因素。结果表明:越冬期灰鹤的主要行为是觅食(77.45%)、警戒(11.17%)和修整(7.82%);行为模式以取食-警戒-取食或取食-修整-取食为主;幼鹤取食行为的时间分配和持续时间均极显著高于成鹤(P0.01),推测幼鹤取食效率低,单次取食持续时间和总时间较长;成鹤警戒行为(P0.01)极显著高于幼鹤,成、幼鹤警戒行为的持续时间无显著差异,取食间隔的警戒频次和时间分配均显著高于幼鹤(P0.05),说明成鹤是通过增加警戒次数来确保幼鹤的安全;取食行为的持续时间和时间分配在整个越冬期均显著升高,取食间隔期间的行走行为也逐渐上升,这与食物资源的可获得性有关;建议适当降低家鸭放牧等人类活动,减少灰鹤越冬期的取食难度;灰鹤不同家庭群的行走行为和取食间隔的行走行为均有显著性差异,推测与各家庭群占有的食物资源质量相关;行为节律上,灰鹤各时段取食行为占总行为的比例均较高,在10:00—10:59、12:00—12:59和16:00—16:59出现取食小高峰;警戒行为保持在一个稳定水平,修整行为高峰出现在取食行为高峰之后,这是因为灰鹤保持取食行为积累疲劳后进行修整;成鹤的取食行为节律性较幼鹤明显,幼鹤的取食行为曲线随机性强,这与幼鹤取食经验不足有关;幼鹤警戒行为的高峰出现在成鹤警戒的低谷,推测与保持整个家庭群较高的总体警戒水平有关。     

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。     

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。     

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。     

云南大山包黑颈鹤日间越冬时间分配和活动节律

2006年10月—2007年4月采用瞬时扫描取样法对云南省昭通市大山包黑颈鹤国家级自然保护区内越冬的黑颈鹤(Grus nigricollis)种群进行了越冬时间分配和日间活动节律的研究。取食是黑颈鹤越冬期间最主要的行为,占(53.05±4.93)%,其次为警戒(18.75±2.65)%、搜寻(10.38±1.34)%和休整(10.32±4.93)%,其余行为依次为:行走(4.90±1.59)%、飞行(1.70±0.38)%、其他(0.55±0.41)%、争斗(0.36±0.25)%。黑颈鹤的出飞时间与日出时间、飞回时间与日落时间分别具有显著的正相关和负相关关系。越冬期间黑颈鹤日间各时间段(1h)(09:00—18:00)和各月份间(2006.11—2007.03)行为差异极显著(P<0.05),并呈现出上午和下午取食双峰、中午休息高峰的规律性变化。气温高时黑颈鹤增加休整时间减少取食时间;气温低时则减少休整时间增加取食时间。偏相关分析说明:环境温度独立地对越冬时间分配产生影响,湿度通过温度对黑颈鹤越冬时间分配产生相反的影响。     

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.     

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.     

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.     

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.     

无人机调查白鹤种群数量和越冬分布

白鹤(Leucogeranus Leucogeranus)是国家Ⅰ级重点保护野生动物,唯一一种极度濒危(CR)鹤类,分布和种群数量是评估其濒危程度的重要参数。无人机调查作为目前生态学研究的一个重要调查方法,为查明白鹤的越冬分布、种群数量及幼鸟比例,于2022年1月使用地面监测结合无人机调查的方法在江西、山东、安徽、湖南、湖北等地的湖泊和农田中开展越冬白鹤调查。野外调查共记录白鹤5607只,网络信息检索在调查区域外的8个地点记录白鹤9只,合计记录越冬白鹤5616只。其中江西鄱阳湖记录到越冬白鹤4813只,占总数的85.7%,主要分布在康山垦殖场、五星垦殖场和成新垦殖场;山东黄河三角洲记录到越冬白鹤625只,占总数的11.1%。安徽、湖南、湖北分别记录到越冬白鹤34,63和72只。对部分群体的白鹤幼鸟数量进行统计,4680只白鹤中,记录到幼鸟674只,幼鸟比例为14.4%,其中湖南越冬白鹤幼鸟比例最高,达28.6%,山东越冬白鹤幼鸟比例最低,为11.5%。调查刷新了白鹤种群数量,证实了山东黄河三角洲是目前除江西鄱阳湖外最重要的白鹤越冬地,缓解了白鹤越冬期过度集中的压力。基于本研究结果,我们...     

甘肃盐池湾黑颈鹤筑巢栖息地偏好及人为干扰的影响

作为高原旗舰物种, 黑颈鹤(Grus nigricolli)是反映高原生态健康状况的重要参考。为了解黑颈鹤如何在多因素作用下适应人类改造过的栖息地环境, 本研究利用遥感解译、最近邻分析与随机森林模型对繁殖于甘肃盐池湾国家级自然保护区党河湿地的黑颈鹤的筑巢栖息地偏好及人为干扰进行研究。2019年和2020年每年的4-9月在党河湿地内对巢位点等数据进行收集。研究结果表明: 党河湿地内巢址与人为干扰的分布位置明显不同, 两者的主要分布区呈现镶嵌状。距深水沼泽距离、距浅水沼泽距离与距湖泊距离是影响黑颈鹤筑巢栖息地选择最关键的3个环境因子。黑颈鹤筑巢时偏好在距离深水沼泽< 125 m、距离浅水沼泽< 130 m、距离湖泊< 270 m的区域内筑巢。黑颈鹤对沼泽、湖泊等资源的偏好是其巢址分布格局的主要驱动力, 而房屋与公路等人为干扰对栖息地选择的影响很小。黑颈鹤筑巢时强烈偏好的栖息地在湿地内占比低、分布聚集且适宜范围有限, 繁殖区域较狭窄。黑颈鹤巢址间距离的上升表明黑颈鹤栖息地质量可能有所下降, 牲畜数量的增长、冬季牧场的多季节利用以及气候变化可能是主要原因。建议在党河湿地内不新增房屋及公路等人为干扰, 同时继续适当限牧, 并给予牧民充足的经济补偿。     

Habitat but not group size or recent predator activity affect corvid collective vigilance at carcasses

Vigilance is an important anti-predator behaviour that can be an indicator of the predation risk faced by potential prey animals. Here, we assess the collective vigilance, or the vigilance level of an entire group, of corvids (Family: Corvidae) at experimentally placed carcasses in a desert environment in Australia. Specifically, we explore the relationship between collective vigilance levels and the habitat in which the carcass was placed, the time since a potential predator (dingo Canis dingo, wedge-tailed eagle Aquila audax or red fox Vulpes vulpes) was present at a carcass, and the group size of corvids around the carcass. We found that corvids are more vigilant in open habitat, but that group size and the recent presence of a potential predator does not affect the collective vigilance behaviour of corvids. The results demonstrate the important link between habitat and vigilance, and that animals may adopt anti-predator behaviours irrespective of the size of the group in which they occur or the recent presence of a potential predator.     

Vigilance and grouping in the southern African ground squirrel (Xerus inauris)

Animals may form groups in response to the foraging–vigilance trade‐off, through enhanced predator detection (collective detection hypothesis) or reduced predation risk to the individual (dilution hypothesis), allowing individuals to decrease vigilance levels. Both hypotheses predict decreasing individual vigilance levels with increasing group size; however, the collective detection hypothesis also predicts increasing overall group vigilance with increasing group size. However, in species in which vigilance and foraging are not mutually exclusive, where vigilance may not be as costly, neither of these hypotheses may apply. Here, we examine the relationship between group size and vigilance in the social Cape ground squirrel (Xerus inauris), a species that can combine foraging and vigilance behaviours. Ten groups were observed using scan sampling, measuring both group and individual vigilance and group size. A negative relationship existed between individual vigilance and group size and a positive relationship between group vigilance and group size. Therefore, in Cape ground squirrels, vigilance seems to be costly even though it can be combined with foraging behaviours. Furthermore, group vigilance behaviour gives support to the collective detection hypothesis, whilst individual vigilance gives support to both hypotheses.     

Common cranesGrus grus and habitat management in holm oak dehesas of Spain

Changes in agricultural policies have favored tree clearing and removal of shrubs to favor intensive farming in the holm oakQuercus ilex dehesas of Iberia. The effect of these changes for bird species wintering in this agricultural landscape has been rarely analyzed. Here the effects are studied of farming changes in traditional holm oak dehesas on population size, social structure and time budget of common cranesGrus grus in eight traditional wintering sites in Spain. The role of acorn availability, the principal winter food of the species, in relation to management is also assessed. The number of cranes in each wintering area was not related to degree of agricultural intensification, nor to the availability of acorns. However, crane flock size decreased, and the relative occurrence of isolated family groups increased in less intensively transformed wintering areas characterized by the predominance of fallow lands. The age composition of crane flocks varied with acorn availability since more juveniles were reported in areas with relatively lower acorn availability. Time budget was also related to acorn availability, since cranes spent relatively more time preening in those wintering areas with higher availability of acorns. Globally evidence suggests the need to maintain the fallow land (posíos) to sustain the wintering populations of cranes in Iberia, since this agro-grazing system maintains most of the juvenile cranes using the western migratory route.     

若尔盖湿地保护区黑颈鹤巢期及影响因子

对鸟类巢期的研究不仅可以丰富鸟类繁殖生物学资料,也可为理解和研究鸟类的生态适应与进化提供重要线索和依据。2013—2014年3月份至7月份,在若尔盖湿地保护区及周边对55对繁殖黑颈鹤的营巢时长进行了研究,调查结果显示:黑颈鹤的营巢时长在0.5—40 d之间,平均巢期为(6.7±9.3)d;在其营造的4种巢型中,巢期长短依次为:泥堆巢草堆巢草墩巢岛地巢,且差异性极显著(P0.001),巢期与巢型显著相关(r=0.728);在其营巢的3种巢址生境中,巢期长短依次为:湖泊生境沼泽生境河流生境,且差异性极显著(P0.001),巢期与巢址生境显著相关(r=0.315);从不同营巢月份看,巢期长短依次为:4月份巢5月份巢6月份巢,且差异性极显著(P0.01),巢期与筑巢月份显著相关(r=0.664);巢期与巢体积大小具有显著相关性(r=0.856),即营巢时间越长巢体积越大。黑颈鹤的营巢时间长短主要受营巢生境、月份和做巢类型的影响。