地表覆盖物对高原鼠兔栖息地利用的影响

对地表覆盖物对高原鼠兔栖息地利用的作用进行了研究, 地表覆盖物增加后, 高原鼠兔能依据其程度及毗邻生境状况, 或减少地面活动增加对洞道系统的利用; 或转移到较安全区域; 同时摄食行为也发生变化, 显著增加用于防御的时间, 且取食区域几乎集中于洞口附近。研究表明, 高原鼠兔视地表覆盖物为一种捕食风险源, 并对此具有一定的评估能力, 其行为反应实质上是通过对食物获取与风险大小的权衡而做出的一种行为决策。间接捕食风险是高原鼠兔在选择和利用栖息地时所必须考虑的权衡因子, 也是一个重要的摄食代价。     

高寒草甸生态系统中高原鼠兔和高原鼢鼠的捕食风险及生存对策

调查青藏高原高寒草甸生态系统中两种主要啮齿动物及其天敌动物的种群密度,分析天敌动物对两种啮齿类的捕食方式、捕食强度,探讨啮齿类动物的捕食风险及生存对策。研究结果表明,高原鼠兔和高原鼢鼠的种群密度分别为4．97只／hm^2和10．6只／hm^2,而它们的主要天敌赤狐、艾虎和香鼬的种群密度分别为0．16只／100hm^2、0．37只／100hm^2、3只／100hm^2。艾虎和香鼬在取食过程中主要搜寻啮齿类的洞道系统,全部食物几乎都来源于洞道系统内;赤狐或取食地面活动的鼠兔,或挖掘洞口待高原鼢鼠封闭洞口时取食猎物。高原鼠兔在赤狐、艾虎和香鼬的食物中所出现的频次分别为100％、96．1％、100％,高原鼢鼠在3种天敌动物的食物中所出现的频次分别为87．5％、73．2％、0％。3种天敌动物对高原鼠兔和高原鼢鼠的捕食强度分别为0．703％和0．038％,高原鼠兔和高原鼢鼠所承受的捕食风险分别为0．780和0．393。高原鼠兔在高的捕食风险下通过行为对策和繁殖对策增加其适合度,而承受捕食风险较小的高原鼢鼠主要通过封闭的洞道系统和高的存活率增加其适合度。     

捕食风险对高原鼠兔食物大小选择的影响

文章报道了捕食风险条件下高原鼠兔对食物大小选择的格局。在实验箱中放置艾虎以改变捕食风险水平, 食物按体积大小分为4种食物项目, 并测定各项目摄入率和取食单个食物项目的进食时间, 结果表明, 摄入率与进食时间依食物项目体积的增大而增加。将大食物与小食物项目配对并供高原鼠兔选择时, 食物项目的利用率视环境状况而不同。捕食风险处理中, 小食物利用率依其进食时间的减少而增加, 其程度与所配对的食物项目的摄入率和进食时间有关。在捕食风险的作用下, 高原鼠兔的食物选择格局反映了能量摄取与风险避免间的权衡。     

捕食风险对高原鼠兔行为的影响

我们在野外条件下利用赤狐的粪尿增加高原鼠兔的捕食风险 ,采用目标动物抽样法对高原鼠兔的 5种行为进行观察 ,分析天敌动物气味增加与天敌动物数量增加对高原鼠兔行为影响的差异 ,探讨高原鼠兔对捕食风险的权衡能力及面对不同风险的行为决策。 2 0 0 1年 ,在捕食风险处理样方中观察雄性成体 1 8只 ,雌性成体2 5只 ,雄性幼体 35只和雌性幼体 4 2只 ,在对照样方中观察雄性成体 1 4只 ,雌性成体 1 5只 ,雄性幼体 2 2只和雌性幼体 1 1只 ;2 0 0 2年 ,在捕食风险处理样方中观察雄性成体 7只 ,雌性成体 1 2只 ,在对照样方中观察雄性成体 8只 ,雌性成体 1 5只。研究结果表明 :增加赤狐的气味后 ,高原鼠兔通过改变行为策略以适应捕食风险的增加。当气味源刚放入后 ,与对照样方比较 ,高原鼠兔明显增加了观察和鸣叫的频次 ,相应减少了取食的频次 ,且随着时间的推移 ,高原鼠兔并未对气味产生适应性。当天敌动物的数量增加后 ,赤狐气味的增加对高原鼠兔行为的影响较小 ,在捕食风险超出高原鼠兔的耐受范围时 ,高原鼠兔扩散。同时 ,高原鼠兔的行为在雌雄之间、成体与幼体之间没有明显的不同 ,雌雄个体、成幼体均采用相同的行为策略减小捕食风险。以上结果表明 :捕食风险明显影响高原鼠兔的行为 ,高原鼠兔能够权衡捕食     

高原鼠兔与达乌尔鼠兔的摄食行为及对栖息地适应性的研究

通过对标志高原鼠兔和达乌尔鼠兔直接观察的方法,对它们的摄食行为及栖息地适应性进行了研究。两种鼠兔的摄食行为链存在着明显趋同的行为程序时间系列。在摄食活动中,前者用于防御敌害的时间分配较多。而达乌尔鼠兔则花费更多的时间于采食。对小生境内的高大植株进行刈割,是高原鼠兔保持其防御视野开阔,降低被捕食风险的适应性策略;达乌尔鼠兔则善于利用栖息地内高大植株覆盖物作为它们的临时隐蔽所,以有效地躲避敌害。这表明它们具有反捕食的行为调控能力,也证明捕食风险强化了物种对栖息地的选择。     

赤狐气味对高原鼠兔繁殖的影响

在野外条件下,利用赤狐的粪尿气味增加高原鼠兔的捕食风险,研究捕食风险对高原鼠兔繁殖的影响。结果表明,作为衡量高原鼠兔繁殖投入大小的定量指标,成体高原鼠兔的体重变化在捕食风险处理样方与对照样方之间没有明显的不同,随着繁殖期的延长,两样方内雌雄个体的体重均显减少,说明捕食风险对高原鼠兔的繁殖投入无明显影响,因此,捕食风险对幼体的生长、发育也无明显的作用。捕食风险增加后,高原鼠兔平均每个雌性成体拥有的后代数目、性比和居留率与对照样方比较均无明显不同,但是由于扩散等原因使每个雄性成体拥有的后代数、繁殖期结束后幼体的性比有明显的差异。以上结果并未显示出捕食气味作为捕食风险对高原鼠兔的繁殖产生抑制作用,其主要原因是捕食风险的类型不同和研究期间高原鼠兔本身承受的捕食风险较大,高原鼠兔可能通过行为变化调节捕食风险增加对其产生的不利影响。     

内蒙古达乌尔鼠兔越冬群体大小 与捕食风险的研究

达乌尔鼠兔(Ochotona dauurica)是内蒙古草原的主要鼠种之一,分布在草地和榆树(Ulmus pumila)疏林下,在冬季会形成数量不同的越冬集群。达乌尔鼠兔在秋季和冬季常常会受到艾虎(Mustela eversmanii)的攻击。为研究达乌尔鼠兔越冬集群大小与来自艾虎的捕食风险的关系,2004年10月上旬在内蒙古锡林郭勒盟锡林浩特市白音锡勒牧场的五连榆树疏林地,以洞口计数法调查分布于林下的达乌尔鼠兔种群,采用洞口数量多少作为鼠兔秋季集群大小的指标。清除实验区达乌尔鼠兔洞群范围内的所有鼬科动物粪便。1个月之后,分别统计洞群留有艾虎粪便和洞群被艾虎挖掘过的达乌尔鼠兔家族数量,分析了艾虎对不同大小集群的达乌尔鼠兔家群的捕食选择偏好。统计结果显示,艾虎在集群大的达乌尔鼠兔家群周围更容易留下粪便(P 0.05),并且更倾向挖掘集群大的达乌尔鼠兔家群(P 0.05)。这表明达乌尔鼠兔越冬家群成员数量越多,其来自艾虎的捕食风险就越大。从本次实验结果来看,越冬期间天敌选择性的捕食风险的存在抑制了达乌尔鼠兔越冬集群的增长,从而使其越冬家族数量维持在限定的规模和水平。     

高原鼠兔对垂穗披碱草及豆科植物生长特性的影响

高原鼠兔通过取食对高寒草甸产生广泛的影响,从而影响高寒草甸植物的个体生长特征。2012年在青海省果洛州矮嵩草草甸,以高原鼠兔喜食植物(豆科和垂穗披碱草)为研究对象,通过小区控制研究高原鼠兔取食（对照组、低密度处理组、中密度处理组、高密度处理组）对植物个体生长特征的影响。研究显示：在实验前期（5月和6月）,喜食植物个体特征基本上无明显规律;在实验后期（7月和8月）,随着高原鼠兔取食强度增大,高原鼠兔喜食植物的地上生物量、平均高度、分蘖枝数、分蘖枝长度、丛幅均呈下降趋势;叶片长度和叶片数均呈先上升后下降态势。2.高原鼠兔喜食植物个体特征的增长率对照组显著高于高密度处理组（P<0.05）。以上结果表明(1)高寒草甸豆类植物在其生长后期与高原鼠兔的取食强度呈一定的负相关关系;高原鼠兔取食强度在阈值（中、低密度）内有利于垂穗披碱草的生长,但超过其阈值的取食强度不利于其生长。(2)高原鼠兔高强度的取食对其喜食植物个体的生长率有明显的抑制作用。     

大鵟对不同数量布氏田鼠越冬洞群的选择偏好

布氏田鼠(Lasiopodomys brandtii)在冬季会形成数量不同的越冬集群,不同大小的集群会面临着不同的捕食风险。2004年秋季在内蒙古锡林郭勒盟典型鼠害草场,采用鼠洞口数量多少作为布氏田鼠秋季集群大小的指标,分析了大鵟(Buteo hemilasius)对不同大小集群的布氏田鼠集群的捕食选择偏好。结果表明：大鵟优先选择集群数量高的布氏田鼠集群,这表明数量高的布氏田鼠集群更容易吸引猛禽类天敌的捕食,该结果可以解释布氏田鼠在秋季出现的分群行为。由于捕食风险的存在,布氏田鼠借助秋季的分群行为以降低冬季集群大小和被捕食风险,从而避免成为捕食性天敌首要攻击的目标,该结果验证了最优集群理论。     

高寒地区植食性小哺乳动物的越冬对策

利用多年工作积累的观察资料 ,讨论几种植食性小哺乳动物的越冬对策。其中 ,高原鼢鼠、甘肃鼠兔和根田鼠均贮存食物 ,以减少寒冷条件下的取食暴露。高原鼢鼠以个体为单位贮存和利用贮存食物 ,相互之间不协作 ;而两种地面活动的种类则可能以家庭为单位贮存和分享越冬食物。喜马拉雅旱獭体型较大 ,不贮存食物 ,它以冬眠方式越冬 ,这是一种对食物依赖最小的方式。高原鼠兔 ,既不贮存食物 ,也不进入冬眠 ,而是主要靠增加身体产热能力来保持体温 ,抵御严寒。作者认为 ,动物自身的生理限制、生活方式、环境条件以及捕食风险等诸多因素的综合作用决定动物的越冬对策。     

集群数量和采食距离对储草期布氏田鼠警戒频次的影响

警戒行为是动物对环境中潜在危险做出的反应。为探究影响警戒行为的因素,作者在内蒙古锡林郭勒典型草原区进行了集群数量和采食距离对储草期自由生活状态下布氏田鼠(Lasiopodomys brandtii)警戒频次影响的实验。通过标志重捕法和人工去除法设定3个集群数量梯度(11只、6只和3只),利用人工食物站设置4个采食距离梯度(5 m、10 m、20 m、30 m),共12个梯度组合,每个组合分别观察60次采食过程,共计观察720次,并记录采食过程中的警戒频次。对数据进行双因素方差分析,集群规模和采食距离对布氏田鼠警戒行为频次均有极显著影响(P0.01)。随着集群数量减小,布氏田鼠单次采食的警戒频次显著增加;采食距离越远,单次采食的警戒频次越高;集群数量和采食距离之间还存在极显著的交互作用(P0.01)。集群数量增大意味着采食距离的增加,为保证储草效率,布氏田鼠种群会在增加集群数量和减小采食距离之间权衡,最终集群数量维持在中等水平,支持了最优集群理论。     

Patch use, apprehension, and vigilance behavior of Nubian Ibex under perceived risk of predation

Foraging theory predicts that animals will sacrifice feedingeffort in order to reduce predation risk. Once a forager choosesa habitat, it must decide how to allocate its foraging effort.Nubian Ibex are diurnal, social, cliff-dwelling herbivores.Many of their characteristics seem to have evolved as responsesto predation risk. In order to assess the effects that perceivedrisk of predation might have on foraging behavior of free-rangingNubian Ibex in the Negev Desert, Israel, we measured giving-updensities (GUDs) in artificial food patches and used them togauge apprehension level. (Apprehension can be defined as areduction in attention devoted to performing an activity asa consequence of reallocating attention to detecting or respondingto predation risk. A forager can also be vigilant. Vigilanceis often defined as time spent scanning the surroundings withthe head up.) We also quantified time budgeting using focalobservation of individual Nubian Ibex. Habitat preferences andpatch selectivity as a measure of apprehension were considered.In particular, we tested the effect of distance from refugeon GUDs, the effect of micropatch structure on selectivity,and the effect of distance from the refuge and group size onNubian Ibex vigilance level and apprehension. Nubian Ibex allocatetheir foraging effort more toward patches closer to the escapeterrain. At the same time, Nubian Ibex are more apprehensiveat intermediate distances from the cliff edge than nearer thecliff, and their use of vigilance increases with distance fromthe cliff edge. These results suggest that Nubian Ibex may switchfrom apprehension to a more extreme behavior of vigilance atgreater distances from the refuge. This study demonstrated theuse of antipredatory behaviors, apprehension, and vigilanceby a forager. Estimating apprehension and vigilance levels ofa forager simultaneously gives a more complete and accuratepicture of how the habitat is perceived by them and combinedwith measurements of GUD allow a more accurate assessment ofhabitat quality.     

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.     

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.     

Foraging Patch Selection in Winter: A Balance between Predation Risk and Thermoregulation Benefit

In winter, foraging activity is intended to optimize food search while minimizing both thermoregulation costs and predation risk. Here we quantify the relative importance of thermoregulation and predation in foraging patch selection of woodland birds wintering in a Mediterranean montane forest. Specifically, we account for thermoregulation benefits related to temperature, and predation risk associated with both illumination of the feeding patch and distance to the nearest refuge provided by vegetation. We measured the amount of time that 38 marked individual birds belonging to five small passerine species spent foraging at artificial feeders. Feeders were located in forest patches that vary in distance to protective cover and exposure to sun radiation; temperature and illumination were registered locally by data loggers. Our results support the influence of both thermoregulation benefits and predation costs on feeding patch choice. The influence of distance to refuge (negative relationship) was nearly three times higher than that of temperature (positive relationship) in determining total foraging time spent at a patch. Light intensity had a negligible and no significant effect. This pattern was generalizable among species and individuals within species, and highlights the preponderance of latent predation risk over thermoregulation benefits on foraging decisions of birds wintering in temperate Mediterranean forests.     

A comparative analysis of vigilance in birds

Individual variation in vigilance is known to vary with factors such as group size but the ecological determinants of vigilance among species have not been examined thus far in a systematic fashion. Earlier analyses suggested that vigilance should be lower in larger species and in species living in larger groups. These analyses were based on a small number of species and failed to take into account phylogenetic relationships among species. Here, I examined ecological determinants of vigilance in a large sample of bird species using a phylogenetic framework. I focused on vigilance in foraging groups of birds in the non-breeding season. Among species, vigilance by solitary foragers was not influenced by body mass. However, among species, asymptotic vigilance, the plateau reached by vigilance in larger groups, decreased with increasing group size in vegetarian clades but not in carnivorous clades. Asymptotic vigilance also increased with increasing body mass in vegetarian clades but not in carnivorous clades. Increasing group size may allow species to reduce vigilance in response to decreased predation risk. Increasing body mass may allow species to increase vigilance because more non-foraging time is available in larger species. Diet may modulate the effect of body mass and group size through factors such as within-group vigilance or foraging techniques.     

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.     

Detecting predators and locating competitors while foraging: an experimental study of a medium-sized herbivore in an African savanna

Vigilance allows individuals to escape from predators, but it also reduces time for other activities which determine fitness, in particular resource acquisition. The principles determining how prey trade time between the detection of predators and food acquisition are not fully understood, particularly in herbivores because of many potential confounding factors (such as group size), and the ability of these animals to be vigilant while handling food. We designed a fertilization experiment to manipulate the quality of resources, and compared awareness (distinguishing apprehensive foraging and vigilance) of wild impalas (Aepyceros melampus) foraging on patches of different grass height and quality in a wilderness area with a full community of predators. While handling food, these animals can allocate time to other functions. The impalas were aware of their environment less often when on good food patches and when the grass was short. The animals spent more time in apprehensive foraging when grass was tall, and no other variable affected apprehensive behavior. The probability of exhibiting a vigilance posture decreased with group size. The interaction between grass height and patch enrichment also affected the time spent in vigilance, suggesting that resource quality was the main driver when visibility is good, and the risk of predation the main driver when the risk is high. We discuss various possible mechanisms underlying the perception of predation risk: foraging strategy, opportunities for scrounging, and inter-individual interference. Overall, this experiment shows that improving patch quality modifies the trade-off between vigilance and foraging in favor of feeding, but vigilance remains ultimately driven by the visibility of predators by foragers within their feeding patches.     

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.