岩松鼠的食物贮藏行为

食物贮藏是许多动物对不可预见的食物供应变化的一种重要适应.岩松鼠(Sciurotamias davidianus)为中国特有物种,广泛分布于华北地区的山地和丘陵地带.作者在北京市东灵山地区建造半自然围栏(4 m×3 m× 1 m),以核桃(Juglans regia)和山杏(Prunus armeniaca)种子为备选食物,对岩松鼠(12只)的食物贮藏行为进行了研究.数据的统计分析采用SPSS for Windows进行.研究结果表明1)岩松鼠表现出集中和分散两种食物贮藏方式,而分散贮藏是其偏好的贮藏方式;2) 当遇到贮藏食物被盗窃时,岩松鼠倾向于搬运更多的食物进行集中和分散贮藏;3)岩松鼠没有在食盘就地取食任何一种食物;把食物搬离后,岩松鼠所取食的山杏种子数量明显多于所取食的核桃数量;4)岩松鼠只选择核桃进行分散贮藏;5)岩松鼠对核桃的搬运距离大于山杏种子.本研究结果提示,在自然条件下,岩松鼠对核桃和山杏的天然更新起着不同的作用[动物学报 51(3)376-382,2005].     

围栏条件下社鼠的食物贮藏行为

在北京东灵山地区, 用活捕笼从野外捕获社鼠, 并在野外建造围栏, 以山杏种子和辽东栎坚果为备选食物, 对围栏条件下社鼠的食物贮藏行为进行了研究。数据的统计分析采用SPSS for Windows (version 1010) 进行。结果表明: 1) 社鼠主要表现出食物的集中贮藏方式; 2) 贮藏食物被模拟盗窃使社鼠的食物贮藏活动加强; 3)社鼠倾向于选择山杏种子进行贮藏, 而对辽东栎坚果则取食较多。文中还讨论了食物类型、模拟盗窃等因素对社鼠食物贮藏行为的影响。     

围栏条件下岩松鼠对核桃的贮藏

许多动物都能够贮藏食物以应对一定时期的食物短缺。岩松鼠是中国特有的一种啮齿动物,主要分布于华北山地和丘陵地区,对于其食物贮藏行为还了解不多。我们在北京东灵山地区建立了半自然围栏,在围栏一角设置一木质巢箱供其休息,在围栏中心设一食盘(给食处),就岩松鼠对核桃(已预先标记)的贮藏特征进行了观察。每只受试动物在实验开始前6 h断食,于13∶ 00 左右,随机选择1 只岩松鼠,将其放入围栏中适应2 ～ 3 h。随后,在给食处放置20 枚已标记的核桃。于第二天同一时间进入围栏,检查并记录岩松鼠对核桃的处理情况,将埋藏、取食和剩余的核桃全部移出,另置20 枚新的核桃,继续实验。第三天,重复上述检查和记录,将岩松鼠及被其处理的核桃全部移出,清理巢箱、整理围栏。随后放入第二只岩松鼠,重复上述实验过程,如遇恶劣气候,则实验顺延,直至实验结束。结果表明:岩松鼠兼具集中贮藏和分散贮藏两种食物贮藏方式,并且分散方式是其主要的食物贮藏方式;岩松鼠在围栏中选择一定的位置贮藏食物;文中还讨论了可能影响岩松鼠食物贮藏的因素。     

围栏条件下中华姬鼠和北社鼠对完好和虫蛀锐齿槲栎种子的取食和贮藏行为差异

正种子贮藏是啮齿动物利用食物资源的一种适应性行为(Vander Wall,1990)。经过长期的进化,啮齿动物形成了两种主要的贮藏食物行为:一是集中贮藏,即把收集的大量食物集中在其洞穴或临时栖息场所贮藏,通常有少量贮藏点,每个贮藏点有大量种子;二是分散贮藏,即把收集的种子分散贮藏在相对较大范围或其巢域周围,通常有大量贮藏点,但每个贮藏点仅贮藏约1～3粒种子。与集中贮藏相比,分散贮藏扩展了植物种子的分布范围,降低了种子被动物取食的概率。由于被分散贮藏的     

Spacing Behavior of a Non‐Larder‐Hoarding Tamiasciurus: A Study of Mearns's Squirrels in Xeric Coniferous Forests

In ecosystems with seasonal fluctuations in food supply many species use two strategies to store food: larder hoarding and scatter hoarding. However, because species at different geographic locations may experience distinct environmental conditions, differences in hoarding behavior may occur. Tree squirrels in the genus Tamiasciurus display variation in hoarding behavior. Whereas red (Tamiasciurus hudsonicus) and Douglas's (Tamiasciurus douglasii) squirrels in mesic coniferous forests defend territories centered around larder hoards maintaining non‐overlapping home ranges, red squirrels in deciduous forests defend small scatter‐hoarded caches of cones maintaining overlapping home ranges. As in other rodent species, variation in hoarding behavior appears to influence the spacing behavior of red and Douglas's squirrels. In contrast, Mearns's squirrels (Tamiasciurus mearnsi) in xeric coniferous forests neither rely on larder hoards nor appear to display territorial behavior. Unfortunately, little is known about the ecology of this southernmost Tamiasciurus. Using radiotelemetry, we estimated home‐range size, overlap, and maximum distance traveled from nest to examine the spacing behavior of Mearns's squirrels. Similar to scatter‐hoarding rodents, maximum distance traveled from nest was greater for males during mating season, whereas those of females were similar year round. Although no seasonal differences were detected, male home ranges were three times larger during mating season, whereas those of females were smaller and displayed a minor variation between seasons. Home ranges were overlapped year round but contrary to our expectations, overlap was greater during mating season for both sexes, with no detectable relationship between male home‐range size and the number of females overlapped during mating season. Overall, the results appear to support our hypothesis that in the absence of larder hoards, the spacing behavior of Mearns's squirrels should be different from larder‐hoarding congeners and more similar to scatter‐hoarding rodents.     

Scatter hoarding by kangaroo rats (Dipodomys merriami) and pilferage from their caches

We observed radio-implanted Merriam's kangaroo rats disposingof 10-g bonanzas of rolled oats in 48 trials in the field. Theprincipal determinant of the initial disposition of discoveredfood was apparently its distance from the day burrow: food foundwithin about 10m was mainly larder hoarded, whereas food encounteredfarther afield was usually dispersed immediately in shallowcaches. Cache sites were newly dug for the purpose and not reused;most caches were nearer the current day burrow than was thefood source, but a few were placed far from both the cacher'sday burrow and its habitual nocturnal range. An experiment withartificial caches indicated that security from discovery increaseswith spacing and with proximity to perennial shrubs. Nine kangaroorats cached dyed food, and fecal dye traces revealed extensivepilferage from five of them, by both conspecifics and otherrodent species. Limited evidence indicates that food encounterednearer home and initially larder hoarded was more secure frompilferage than food initially scattered, and yet kangaroo ratswere observed to scatter caches soon after initial larder hoarding.A kangaroo rat whose dyed stores escaped pilferage fed fromthem at intervals for at least 12 days. Even cachers who incurredpilferage made as much, or more, use of their caches as anythief, suggesting that scattering caches may be a defense againstcatastrophic losses.     

食物源距离对中华姬鼠贮藏策略的影响

在围栏条件下, 检验食物源距离对中华姬鼠贮藏策略的影响,并检验根据快速隔离假说和最优贮藏空间分布模型做出的两个预测：（1）3个释放点无论距离远近,种子均先被大量分散贮藏在各个释放点周围;（2）随着释放点远离巢穴,种子被集中贮藏的比例下降。结果发现: 分散贮藏是中华姬鼠的主要贮藏方式,3个释放点种子被分散贮藏的数量无显著差异,这个结果支持预测（1）;3个种子释放点（1m,5m,13m）被集中贮藏的比例分别为(1m: 6.25%; 5m: 1.88%; 13m: 1.25%),说明被集中贮藏的种子数量随种子释放点与巢穴距离的增大而逐渐减少,这个结果支持预测（2）。本研究表明中华姬鼠应对捕食风险升高的策略是降低取食量和贮藏量。     

Effects of habitat and season on removal and hoarding of seeds of wild apricot (Prunus armeniaca) by small rodents

The wild apricot (Prunus armeniaca) is widely distributed in the Donglingshan Mountains of Mentougou District of Beijing, China, where its seeds may be an important food resource for rodents. Predation, removal and hoarding of seeds by rodents will inevitably affect the spatio-temporal pattern of seed fate of wild apricot in this area. By marking and releasing tagged seeds of wild apricot, we investigated seeds survival, scatter-hoarding, cache size and seedling establishment, and the preference of micro-habitats used by rodents to store seeds. The results showed that: (1) rodents in this area hoarded food intensively in autumn, as well as in spring and summer. (2) There were significant effects of habitat and season on removal rate of tagged seeds at releasing plots. In both two types of habitats, Low and High shrub, tagged seeds were removed most rapidly by rodents in autumn, at intermediate rates in spring and least rapidly in summer. (3) During three seasons, mean dispersal distance of scatter-hoarded seeds in Low shrub habitat was greater than that in High shrub. Most removed seeds were buried within 21.0 m of the releasing plots. (4) In both two types of habitats, Low and High shrub, rodents tended to carry seeds to US (Under shrub) and SE (Shrub edge) microhabitats for scatter-hoarding or predation. (5) Among the caches made by rodents, most caches contained only one seed, but up to three seeds were observed; caches of 2–3 seeds were common in autumn. (6) By comparing dental marks, we determined that large field mice (Apodemus peninsulae) and David’s rock squirrels (Sciurotamias davidianus) contributed to removal and predation of released tagged seeds. However, only the large field mice exerted a pivotal and positive role on the burial of dispersed seeds. (7) Establishment of three seedlings originated from seeds buried by rodents was documented in High shrub habitat.     

Sensitivity to Seed Germination Schedule by Scatter‐Hoarding Pére David's Rock Squirrels During Mast and Non‐Mast Years

Mast seeding by oaks (Quercus) and other seed species has fundamental impacts on the behavior of individual scatter‐hoarding animals as well as their population and community dynamics. Scatter‐hoarding squirrels are highly sensitive to acorn germination schedule and stop germination by removing the embryo of non‐dormant acorns to prevent losses of energy and nutrients by rapid germination. However, we know little about how this behavior is affected by mast seeding. We investigated foraging decisions made by free‐ranging Pére David's rock squirrel (Sciurotamias davidianus) with three species that produce non‐dormant seeds and two species that produce dormant seeds (Fagaceae) during mast and non‐mast years in Central China from 2007–2010. Consistent with the food perishability hypothesis that squirrels make decisions to minimize the perishability of their caches, non‐dormant seeds were hoarded at a lower rate, but had a higher probability of embryo removal than dormant seeds. Compared with non‐mast years, a lower proportion of seeds were harvested during mast years, but a higher proportion of the harvested seeds were hoarded. In addition, the probability of embryo removal in the hoarded seeds was higher during mast years. Squirrels also dispersed seeds at shorter distances during mast years. Moreover, the interactions between mast seeding and seed germination schedule had significant effects on squirrel foraging decisions, including embryo removal. This study is the first to demonstrate that squirrels show greater sensitivity to seed germination potential when seeds are abundant.     

Application of radiotelemetry to the survey of acorn dispersal byApodemus mice

We examined the applicability of radiotelemetry to studies of acorn dispersal byApodemus mice and compared its efficiency with the of this spool-and-line method. Installation of a transmitter (2.2 g) onto acorns did not interfere with the transporting and feeding behavior of the mice. We were able to detect all transmitter-installed acorns and follow the daily changes in the sites in which they were hoarded, while we missed 59% of the spool-tied acorns due to mice breaking the threads. Mice carried transmitter-installed acorns farther than spool-tied ones. The radiotelemetry method is superior to the spool-and-line method and useful for the study of hoarding behavior in rodents.     

Scatter‐hoarding and cache pilfering of rodents in response to seed abundance

Seed caching and reciprocal cache pilferage play an important role in the coexistence of food‐hoarding animals. Understanding what affects seed caching and how cache pilferage occurs is an important question in seed dispersal ecology. However, tracking seed fate and cache pilferage presents substantial practical difficulties. Siberian chipmunks Tamias sibiricus always remove the entire pericarp when scatter‐hoarding acorns of Mongolian oak Quercus mongolica, whereas wood mice Apodemus peninsulae often store whole acorns in their caches. These differences in behavior provide an opportunity to investigate unilateral cache pilferage of T. sibiricus from A. peninsulae in response to seed abundance. In this study, tagged acorns were released at the peak and end periods of seed rain from Q. mongolica. This allowed us to investigate seed caching and unilateral cache pilferage at different seed abundances. We found that a higher proportion of acorns were cached at lower level of seed abundance (toward the end of seed rain), mainly because T. sibiricus rather than A. peninsulae scatter‐hoarded significantly more acorns at this time. Cache distances decreased with increasing seed abundance, indicating that acorns were cached further away and into smaller caches at lower seed abundance. Unexpectedly, unilateral cache pilferage by T. sibiricus was not significantly influenced by seed abundance—remaining at around 28% during both periods of high and low seed abundance.     

Inter-Population Variation in Hoarding Behaviour in Degus, Octodon degus

Although foraging comprises a set of behaviours that typically vary with resource availability and/or climatic conditions, few studies have analysed how foraging, particularly food hoarding, varies across populations inhabiting different habitats. We carried out an inter-population study on foraging behaviour with the caviomorph rodent Octodon degus collected from two geographically separated populations in central Chile, with contrasting climates. One population was located in a mountainous zone (at 2600 m elevation) characterized by a high-altitude climate. The other population was from a low-altitude Mediterranean climate zone (450 m elevation). Under laboratory conditions, we measured population-specific differences in food consumption and hoarding by recording food utilization. We also assessed whether acclimation played a role in behavioural differences, by using two different sets of animals that had been in captivity for (1) 2 wk or (2) 6 mo, under common conditions. The results showed variation in food hoarding between populations. Individuals from the low-altitude population exclusively displayed scatter hoarding behaviour. In contrast, high-altitude animals carried out larder hoarding combined with scatter hoarding (37.4% and 62.6% respectively). There was no intra-population variation between degus with different acclimation periods under captivity, thus inter-population differences in larder hoarding were maintained despite 6 mo of acclimation to a common environment. The geographic variation observed suggests that larder hoarding is favoured under harsher environmental conditions. We discuss some probable causes for this variation. The lack of effect of acclimation suggests that inter-population differences in larder hoarding might be the result of local adaptation or, less likely, it corresponds to an ontogenetically acquired irreversible behaviour.     

啮齿动物的分散贮食行为

食物贮藏是许多动物重要的适应性行为,分散贮藏的食物以植物种子为主。每个贮藏点贮藏数量不等的食物项目。啮齿动物分散贮藏食物之后,可降低食物被其他个体获取的机率,提高对食物资源的控制能力,最终有利于自身的生存和繁殖成功。植物种子被贮藏之后,可减少非贮食鼠类对种子的取食。同时,合适的微生境和埋藏有利于种子萌发、幼苗建成和植物的更新;使植物的分布区得以扩展。探讨啮齿动物的分散贮食行为,能够更好地理解食物贮藏在啮齿动物生活史中的作用,进一步认识鼠类和植物的相互关系以及不同啮齿动物在群落形成中的潜在作用。本综述了啮齿动物分散贮食的研究进展,并提出今后工作中的几点建议。     

鼠类对栓皮栎和锥栗种子的贮藏:验证高单宁假说

种子中的单宁影响鼠类的取食和贮藏策略。高单宁假说认为鼠类倾向于优先贮藏单宁含量高的种子、取食单宁含量低的种子。同域分布的鼠类对种子的取食和贮藏选择是否符合高单宁假说的预测尚缺乏足够的实验证据。本研究通过围栏实验研究了四川都江堰地区亚热带常绿阔叶林中的小泡巨鼠（Leopoldamys edwardsi）、针毛鼠（Niveventer fulvescens）、北社鼠（N.confucianus）、高山姬鼠（Apodemus chevrieri）、中华姬鼠（A.draco）对单宁含量高的栓皮栎（Quercus variabilis）种子和单宁含量低的锥栗（Castanea henryi）种子的取食和贮藏偏好,旨在进一步验证高单宁假说。结果表明：（1）5种鼠均偏好取食锥栗种子,很少取食或不取食栓皮栎种子;（2）仅具集中贮藏习性的针毛鼠和北社鼠偏好集中贮藏锥栗种子;兼具集中和分散贮藏习性的小泡巨鼠和高山姬鼠也倾向于贮藏锥栗种子;（3）兼具集中和分散贮藏习性的中华姬鼠偏好分散贮藏栓皮栎种子,集中贮藏锥栗种子。研究结果并不完全支持高单宁假说,表明同域分布的鼠类对种子取食和贮藏偏好表现出种间分化,高单宁假说在同域分布鼠类中可能不具有普遍性。     

Weak olfaction increases seed scatter‐hoarding by Siberian chipmunks: implication in shaping plant–animal interactions

Scatter‐hoarding of seeds by animals plays an essential role in seed dispersal of plants and in shaping plant–animal interactions in forest ecosystems, but the function of scatter‐hoarding behavior is still unclear. We hypothesize that weak olfactory cues between seeds and scatter‐hoarding animals would increase scatter‐hoarding. Using a rodent–plant system of Siberian chipmunks Tamias sibiricus and Korean pines Pinus koraiensis, we tested the effects on seed scatter‐hoarding intensity by measuring and modifying the seed odor intensities and the abilities of the animals to detect seed odor. Siberian chipmunks seemed to scatter‐hoard more seeds with weaker odor signals, and Siberian chipmunks with reduced olfactory ability scatter‐hoarded more seeds, supporting our hypothesis. Our studies suggest that olfaction may have played an important role in the evolution of the strength of seed odor and scatter‐hoarding behavior of animals, and in shaping plant–animal interactions.     

Pregnancy coloration in macaques may act as a warning signal to reduce antagonism by conspecifics

Co-varying traits in acorns such as seed size and germination schedule are important to influence the behavioural decisions of hoarding rodents. Using acorn pairs from cork oak (Quercus variabilis) (large size and short germination schedules) serrate oak (Q. serrata) (small size and short germination schedule) and qinggang (Cyclobalanopsis glauca) (small size and long germination schedule) with contrasting seed size and germination schedule, we conducted a series of experiments to investigate hoarding preferences in response to seed size and germination schedule by Edward's long-tailed rats (Leopoldamys edwardsi) and South China field mice (Apodemus draco) in semi-natural enclosures. We found that the seed size hypothesis was consistently supported: both rodent species ate more small acorns but hoarded more large ones regardless of germination schedules. However, the germination schedule hypothesis was also supported when similar sized acorns were simultaneously provided, e.g. Q. serrata versus C. glauca or germinating versus non-germinating Q. variabilis. Our results, contrary to the studies from North America, indicate that seed size is more important than germination schedules in determining whether the tested animals eat or hoard a given seed.     

围栏条件下捕食风险对岩松鼠贮藏核桃种子行为的影响

面对捕食风险,贮藏植物种子的啮齿动物如何通过调整其贮藏行为,权衡捕食风险和贮食效益,需要深入研究。2006 年9 月,在北京东灵山地区,以黄鼬作为捕食风险源,在半自然围栏内比较了有无捕食风险源时,岩松鼠贮藏的核桃种子的状态,以及埋藏种子在实验围栏内的分布、微生境和搬运距离,以探讨捕食风险对岩松鼠贮藏种子行为的影响。结果表明,捕食风险源存在时,搬运和埋藏种子的比例都显著减少,弃置地表种子的比例无显著差异;埋藏于高风险区的种子比例减少,中风险区和低风险区的埋藏种子比例增加,但差异都不显著;位于裸地和草下的埋藏种子比例减少,位于墙基的埋藏种子比例增加,但差异都不显著;此外,埋藏种子的搬运距离也没有显著差异。可见,捕食风险源存在时,岩松鼠显著减少了搬运和埋藏种子的比例,但对埋藏种子时的埋藏区域和微生境选择,以及搬运距离无显著影响。差异不显著可能与围栏效应有关。     

灌丛高度对啮齿动物贮藏和扩散辽东栎坚果的影响

本文选择高、矮两类灌丛生境, 于2002 年至2003 年在其中释放标记的辽东栎坚果, 并连续记录释放坚果的命运, 以了解生境差异对啮齿动物搬运、取食和贮藏辽东栎坚果的影响以及啮齿动物对微生境的选择等。研究结果表明, 大林姬鼠等啮齿动物对辽东栎坚果的短期取食非常强烈, 而分散贮藏的量则相对较少; 啮齿动物对辽东栎坚果的搬运距离在矮灌丛生境中显著大于高灌丛生境; 啮齿动物倾向于选择灌丛边缘和灌丛下方取食或贮藏辽东栎坚果, 这类微生境也有利于埋藏坚果的萌发。     

Seed dispersal of the Korean pine,Pinus koraiensis,by the red squirrel,Sciurus vulgaris

Seed hoarding behavior of the red squirrel,Sciurus vulgaris, was studied in relation to the amount of dispersed seeds of the Korean pine,Pinus koraiensis, and the distribution of its seedlings. After removing a cone from a tree, squirrels sat on the ground and ripped off its cone scales before transporting it. A mean of 3.2 seeds were scatter-hoarded per hole. Of 7.7×10⁴ mature seeds produced in a 0.21 ha planted Korean pine forest, 22% were estimated to be directly eaten by four squirrels, 9% were hoarded by them in the pine forest and 65% were cached outside the forest. Squirrels rediscovered hoarded seeds frequently, until the ground was covered with snow, during the period from snow fall until seed germination the next spring, few hoarded seeds were utilized. Korean pine seedlings were found up to 600 m from their mother trees. Scatter-hoarding by squirrels extensively contributes to seed dispersal to places suitable for the regeneration of the Korean pine. The large size of the cone, the absciss-layer at the cone penduncle, the infrequent dehiscence of cone scales, the large and wingless seeds, and the thick seed-coats have probably all been specialized to facilitate utilization by GenusSciurus.     

Effects of burrow condition and seed handling time on hoarding strategies of Edward's long-tailed rat (Leopoldamys edwardsi)

Many hoarding rodents use burrows not only for dwelling and protection from natural enemies, but also for food storage. However, little is known how burrows used by scatter-hoarding animals influence their foraging behaviors. In addition, handling time for a given food item has a fundamental impact on hoarding strategies of these hoarding animals: food items with longer handling time are more likely to be hoarded due to increasing predation risk because the animals spend more time outside their burrows if they consumed such food. By providing with two types of artificial burrows (aboveground vs. underground) and two types of food items (i.e. seeds) with contrasting handling times, we investigated how burrow condition and handling time co-influence hoarding strategies of a key scatter-hoarding rodent, Edward's long-tailed rat (Leopoldamys edwardsi) in large enclosures in southwest China. We found that only a few animals larder-hoarded fewer seeds when only aboveground burrows were available, while over 80% of the animals preferred to use the underground burrows and hoard significantly more seeds in the burrows when both aboveground and underground burrows were provided simultaneously. We also found that seed handling time significantly affected hoarding strategies of the animals: they consumed and/or scatter-hoarded more Camellia oleifera seeds with shorter handling time outside the burrow, but consumed and larder-hoarded more Lithocarpus harlandii seeds with longer handling time in underground burrows. Our study indicates that both burrow types and seed handling time have important impacts on hoarding strategies of scatter-hoarding animals.