Clark's Nutcrackers Nucifraga columbiana Remember the Size of Their Cached Seeds

Clark's nutcrackers ( Nucifraga columbiana ) hide thousands of seeds in subterranean caches that they later recover using spatial information about cache location. In two experiments, we tested whether nutcrackers also remember another type of information regarding their caches – the size of the seeds in each cache. We videotaped birds during cache recovery and then measured their bill gape during probing behaviour as captured on the videotape. In experiment one, six birds each experienced two treatments: one that allowed them to cache and then recover large seeds, and the other, an identical treatment using small seeds. During this experiment, all six birds used a wider gape when attempting to recover seeds during the large-seed treatment than during the small-seed treatment, and gape width was significantly correlated with seed size. During experiment two, we presented birds with both large and small seeds within the same caching session. We also increased the retention interval between caching and recovery. These modifications increased the difficulty of the task. Six of the seven birds used a wider gape during seed recovery when digging for caches that contained large seeds than they did when searching for small seeds. The ability to remember the size of seeds placed within caches may serve to increase the likelihood of speedy and successful recovery. It also allows the birds another level of organization of their food supply. These are the first experiments to suggest that Clark's nutcrackers remember more about their caches than location alone.     

Revisits to emptied cache sites by Clark's nutcrackers (Nucifraga columbiana)

During cache recovery experiments, Clark's nutcrackers (Nucifraga columbiana) often revisit cache sites that were emptied during previous recovery sessions (Kamil & Balda 1985). In the current experiments, nutcrackers were allowed to cache pine seeds and recover them in several recovery sessions. The results of two experiments demonstrate that neither leaving signs of previous digging activity nor leaving many pine seeds in the caches affects the tendency to revisit.     

贮藏点深度、大小及基质含水量对花鼠找寻红松种子的影响

2012年8-10月,在黑龙江省伊春市带岭林业局东方红林场,通过人工围栏控制实验,研究了贮藏点深度、大小及基质含水量对花鼠找寻红松种子的影响。发现：(1) 埋藏深度显著影响花鼠对贮藏点的找寻率,埋藏深度为 1 cm 和 2 cm 的找寻率显著高于 4 cm 和 6 cm。(2) 埋藏点大小对花鼠发现种子也有显著影响,埋藏点越大,花鼠发现贮藏点的比例越高。(3) 高的基质含水量利于花鼠找寻贮藏点。结果表明,围栏条件下食物贮藏点深度、大小及基质含水量的改变能显著影响花鼠对贮藏点的找寻。     

Convenience Stores: Repeated Use of Some Cache Sites by Clark's Nutcrackers (Nucifraga columbiana)

Field studies reveal that Clark's nutcrackers (Nucifraga columbiana) cache thousands of seeds in subterranean caches during the autumn in some years, and laboratory studies have shown that these caches can be recovered with high levels of accuracy. Over the life of a bird, numerous cache recovery cycles may be experienced and many of these may be in the same area. Birds may thus develop strong individual preferences for certain sites. These sites may become more memorable and used preferentially. These sites might also be more vulnerable to theft because of continual reuse. In this study, birds were allowed to cache and recover four times from the same set of potential cache sites in a large experimental room. Three of five birds used some cache sites more often than would be expected by chance. However, these birds did not place more seeds per cache, have higher recovery accuracy, or make more revisits to cache sites previously used. Birds did not create caches earlier in previously used holes until cycle 4, when birds actually cached in used sites significantly sooner than in previously unused ones. Birds neither recovered caches earlier from previously used sites nor made error probes earlier at them. Sites that were repeatedly used for caching were holes that were significantly closer to a centrally located feeder than less frequently used sites. Thus, some birds may develop slight preferences for some locations but these sites are not treated preferentially. Birds appear not to gain the advantages, nor suffer the consequences associated with repeated use of the same sites.     

Substrate type affects caching and pilferage of pine seeds by chipmunks

The abiotic environment often influences the ways in which animalsinteract. By affecting the cues associated with buried seeds,the type of substrate used by seed-caching rodents may alterthe relative probabilities of cache pilferage and cache retrieval.We predicted that, after a wildfire, the presence of ash wouldimpair rodents' ability to smell pine seeds on the forest floor.In a laboratory experiment, we compared the foraging success,caching frequency, and cache recovery of chipmunks (six Tamiasamoenus and six T. quadrimaculatus) in ash versus sand substrates.Initial results supported our hypothesis: chipmunks found only2.3% of 108 caches of Jeffrey pine (Pinus jeffreyi) seeds thatwe buried in ash but found 98% of caches in sand. However, chipmunksmade as many or more of their own caches in ash compared withsand (48% for T. amoenus, 73% for T. quadrimaculatus.) Whenforaging for seeds cached in ash by themselves and by otherindividuals, they found significantly higher proportions oftheir own caches (62%) than of caches made by others (25%).However, when foraging in sand, they found high proportionsboth of their own caches and those of others (86 versus 81%).These results suggest that olfaction is less effective in ashthan in sand, that spatial memory enables chipmunks to recovertheir own caches in ash, and that caching in ash may allow animalsto avoid pilferage of stored food. As chipmunks are importantdispersers of seeds, changes in their foraging patterns or competitiveinteractions after fire could significantly affect pine regeneration.     

Clark's Nutcracker (Aves: Corvidae) Spatial Memory: Interference Effects on Cache Recovery Performance?

Clark's nutcrackers, Nucifraga columbiana, accurately v recover thousands of caches per year in the field. Previous experiments have confirmed that these birds possess excellent, long-lasting spatial-memory capabilities. We tested whether resistance to interference is one of the features of nutcracker spatial memory. Experiment 1 tested retroactive interference. Nutcrackers showed no decrease in accuracy overall but performed relatively poorly in their final recovery session. Interference is unlikely to have caused these sites to be poorly remembered because they had fewer neighbouring cache sites than better-remembered sites. Experiment 2 tested for proactive interference. Interference would have caused the experimental birds to be less accurate than control birds. Instead then were slightly more accurate. In experiment 3, nutcrackers were allowed to repeatedly view their cache sites from a cage between caching and recovery. Nutcrackers were less accurate when recovering from cache sites they had viewed. This effect may be due to changes in motivation. Order of caching had no effect on accuracy but nutcrackers were more accurate when recovering caches from central than from peripheral areas of experimental rooms. In summary, these experiments provide further evidence of the remarkable spatial-memory abilities of Clark's nutcrackers and demonstrate that these birds are highly resistant to interference effects on spatial memory. Comparative tests will be needed to test if specialized food storers are exceptionally resistant to interference in spatial memory.     

Bolus recovery by gray jays: an experimental analysis

Gray jays (Perisoreus canadensis) typically store food boli in various sites on conifers. In a laboratory setting we determined whether gray jays recover stored boli by means of olfaction, trial-and-error search or spatial memory. Using an artificial tree with 52 possible caching sites, caching and/or recovery trials were performed with five captive gray jays for the following experiments: (1) no extra visual cues on tree; (2) extra visual cues (pine foliage) attached to tree; (3) pungent-smelling food hidden by observer; (4) one bird allowed to cache food but caches recovered by a second bird; (5) one bird allowed to observe another bird cache food and later permitted to recover those caches. Results supported the memory hypothesis, but cache site preferences were apparent for individual birds. To control for this, an additional experiment (6), in which cache site access was limited by the investigators, was conducted with two new birds. These results also indicated that gray jays use spatial memory to recover stored boli.     

The influence of environmental conditions on cache recovery and cache pilferage by yellow pine chipmunks (Tamias amoenus) and deer mice (Peromyscus maniculatus)

I conducted a field experiment in 10 x 10 m enclosures to explorehow seed and soil moisture levels influence the ability ofknowledgeable and naive rodents to find natural caches of Jeffreypine (Pinus jeffreyi) seeds. Subjects were yellow pine chipmunks(Tamias amoenus) and deer mice (Peromyscus maniculatus) searchingfor caches that they had made, caches made by other individualsof the same species, or caches made by individuals of the otherspecies. Subjects that made caches (knowledgeable subjects)relied on spatial memory to find many of their own caches during recovery sessions, and their ability to locate caches was notaffected by water content of seeds or soil. Naive subjectsfound few caches under dry conditions, but under wet conditions,they located as many caches as did the rodents that made them.Naive subjects apparently relied on olfaction to find caches,a sensory modality that works more effectively under moist conditions. Subjects had as much success foraging for caches made by membersof their own species as for caches made by the other species.I present a hypothesis that predicts how foragers could modifypredominately memory-based search to predominately olfactory-basedsearch as the weather changes from dry to wet. When foragersrely on spatial memory, those foragers find only their own caches, but when they can also use olfaction, they pilfer cachesmade by other individuals. Consequently, the nature of competitiveinteractions among members of the seed-caching guild may changeas the weather changes.     

Food storage by black-capped chickadees: Memory for the location and contents of caches

Black-capped chickadees (Parus atricapillus) store food in a scattered distribution in their winter home range. Several hundred food items may be stored in a day, each in a separate cache site. Previous studies of marsh tits (Parus palustris) and nutcrackers (Nucifraga spp.) have shown that spatial memory is used to relocate caches. Memory for storage sites, if used by black-capped chickadees, is predicted to have four properties. Birds should be able to: (1) accurately relocate cache sites, (2) recall which caches they have previously emptied, (3) recall which sites they have discovered empty (as a result of loss to other animals) and (4) recall what type of food is stored at a cache site. Laboratory experiments show that chickadees do incorporate these kinds of information in memory for cache sites.     

埋藏点深度、间距及大小对花鼠发现向日葵种子的影响

2005 年5 ～6 月,在北京东灵山地区,通过在实验围栏内设置不同深度、间距和大小的埋藏点,研究了花鼠对向日葵种子的发现率。结果表明,花鼠发现埋藏种子的比例随埋藏深度的增加而减少,埋藏深度> 5 cm后,发现比例较低,且不再有明显变化;花鼠发现埋藏种子的比例随埋藏点间距的减小而增加,埋藏点间距<0.5 m后,发现比例较高,且不再有明显变化;花鼠发现埋藏种子的比例随着埋藏点增大而增加,埋藏点种子> 5 g后,发现比例较高,且不再有明显变化。     

Placement, Retrieval, and Memory of Caches by Thirteen-Lined Ground Squirrels

Mechanisms governing placement and retrieval of scatter hoards were investigated in 13-lined ground squirrels ( Spermophilus tridecemlineatus ). The squirrels used several placement tactics, including relatively deep burial of seeds, avoidance of prominent objects, and cryptic placement, which would reduce chance discovery by competitors but that might also increase the difficulty of retrieval for the hoarder. Nevertheless, scatter hoards were unfailingly retrieved within a day or two after placement, despite experimental elimination or displacement of local sensory cues emanating from the sites. Artificial caches placed close to true caches were not discovered, indicating that recovery attempts were quite precise. These results imply that 13-lined ground squirrels rely heavily on spatial memory for retrieval and are the first experimental demonstration of the importance of memory for cache recovery in a natural population of mammals.     

Seed-caching responses to substrate and rock cover by two <Emphasis Type="Italic">Peromyscus</Emphasis> species: implications for pinyon pine establishment

We examined whether pinyon mice (Peromyscus truei) and brush mice (P. boylii) could act as directed dispersal agents of pinyon pine (Pinus edulis) through differential responses to soil particle size and rock cover. In field experiments, we allowed mice to either cache pinyon seeds or recover artificially cached seeds (pilfer) from quadrats containing large- or small-particle soils. Both species placed most (70%) seed caches in small-particle soil. Pilfering was the same from both particle sizes in the first year, while more seeds were pilfered from large-particle soils in the second year. In separate experiments, rock cover interacted with soil particle size, with both species placing over 50% of their caches in small-particle soil with rock cover. Overall, we found greater seed-caching in small-particle soils near rocks, with equal or lower pilfering from small-particle soils, suggesting more seeds would survive in small-particle soils near rock cover. Three lines of evidence supported the hypothesis that mice could act as directed dispersers by moving pinyon seeds to beneficial microsites for germination and establishment. First, in greenhouse experiments, pinyon seed germination was 4 times greater in small-particle soil cores than in large-particle soil cores. Second, soils near rocks had significantly higher water content than areas of open soil at the driest time of the year, a critical factor for seedling survival in the arid southwestern USA. Third, 75% of juvenile pinyon trees were growing in small-particle soils, and 45% were growing near rock nurses.     

Observational visuospatial encoding of the cache locations of others by western scrub-jays (<Emphasis Type="Italic">Aphelocoma californica</Emphasis>)

Western scrub-jays (Aphelocoma californica) hide food and rely on spatial memory to recover their caches at a later date. They also rely on observational spatial memory to steal caches made by other individuals. Successful pilfering may require an understanding of allocentric space because the observer will often be in a different position from the demonstrator when the caching event occurs. We compared cache recovery accuracy of pairs of observers that watched a demonstrator cache food. The pattern of recovery searches showed that observers were more accurate when they had observed the caching event from the same viewing direction as the demonstrator than when they had watched from the opposite direction. Search accuracy was not affected by whether or not the tray-specific local cues provided left–right landmark information (i.e. heterogeneous vs. homogeneous local cues), or whether or not the caching tray location was rotated. Taken together, these results suggest that observers have excellent spatial memory and that they have little difficulty with mental rotation.     

围栏条件下花鼠找寻种子的途径和方式

2009年8月和2010年8月,在黑龙江省带岭区东方红林场半天然围栏内研究了花鼠(Eutamias sibiricus)对地表放置和埋藏种子的发现过程,摸清围栏条件下花鼠找寻种子信号的途径和方式.实验步骤如下:第1天在花鼠巢区对角每释放点地表放置1粒标记种子,共30粒;第2天在巢区对角每释放点埋藏1粒标记种子,共30粒...     

贮藏种子的啮齿动物对油茶种子沉积形式的影响

种子沉积的质量常常涉及种子扩散作用者(如鸟类和小型啮齿动物)对生境和微生境的选择以及种子沉积的形式(如种子埋藏).然而,很少的研究涉及到种子在离开母树后被这些动物沉积在何处.在四川省都江堰一个实验林场的2个林分(次生林和原生林)内,通过追踪用带编号的金属薄片标记的油茶(Camellia oleifera Abel.)种子的命运,研究了贮藏种子的啮齿动物对种子沉积的影响.研究发现:在2个林分内,80%以上的种子被很好地埋藏在0～60 mm深的土壤中,而小部分种子则被放置在地表(但有少量落叶遮盖).小型啮齿动物喜好在灌丛下或灌丛边缘贮藏和取食种子,可能是在这样的微生境下它们在觅食时将遭遇较小的捕食风险.研究还发现,贮藏点的微生境分布随贮藏点等级而逐渐变化:在两个林分内较高等级的贮藏点(如次贮藏点和三级贮藏点)比初级贮藏点有更多的种子被贮藏在灌丛下或灌丛边缘.这表明,啮齿动物对油茶种子的埋藏可能更有益于种子的存活、萌发以及幼苗的建成.啮齿动物将散落在母树下或其附近的油茶种子扩散到不同的微生境,这可能有利于他们遭遇到更多的适宜环境而萌发,实现幼苗补充.     

贮藏种子的啮齿动物对油茶种子沉积形式的影响

种子沉积的质量常常涉及种子扩散作用者(如鸟类和小型啮齿动物)对生境和微生境的选择以及种子沉积的形式(如种子埋藏)。然而,很少的研究涉及到种子在离开母树后被这些动物沉积在何处。在四川省都江堰一个实验林场的2个林分(次生林和原生林)内,通过追踪用带编号的金属薄片标记的油茶(CamelliaoleiferaAbel.)种子的命运,研究了贮藏种子的啮齿动物对种子沉积的影响。研究发现在2个林分内,80%以上的种子被很好地埋藏在0~60mm深的土壤中,而小部分种子则被放置在地表(但有少量落叶遮盖)。小型啮齿动物喜好在灌丛下或灌丛边缘贮藏和取食种子,可能是在这样的微生境下它们在觅食时将遭遇较小的捕食风险。研究还发现,贮藏点的微生境分布随贮藏点等级而逐渐变化在两个林分内较高等级的贮藏点(如次贮藏点和三级贮藏点)比初级贮藏点有更多的种子被贮藏在灌丛下或灌丛边缘。这表明,啮齿动物对油茶种子的埋藏可能更有益于种子的存活、萌发以及幼苗的建成。啮齿动物将散落在母树下或其附近的油茶种子扩散到不同的微生境,这可能有利于他们遭遇到更多的适宜环境而萌发,实现幼苗补充。     

The effects of cache loss on choice of cache sites in black-capped chickadees

Food-storing birds lose a great deal of their stored food toother animals. We examined whether blackcapped chickadees (Parusairicapillus) modify their choice of cache sites using informationthat predicts cache loss. In experiment 1, birds learned toavoid caching at spatial locations where cache loss had previouslyoccurred, but they did not avoid caching near local color cuesthat predicted cache loss. Birds did not modify their generaluse of space in the aviary. Birds also learned to reduce searchingfor caches where spatial location predicted cache loss. Experiment2 confirmed the birds’ ability to discriminate among thespatial locations and the local color cues used in experiment1. In experiment 3, learning a food-rewarded approach to potentialcache sites occurred without any change in the choice of sitesfor caching. We discuss how chickadees selectively associatethe choice of cache site with its consequences, even over delaysof several hours between caching and cache recovery.     

Recaching Decisions of Florida Scrub‐Jays are Sensitive to Ecological Conditions

Food caching animals depend on their caches at times of low food availability. Because stored food is susceptible to being stolen or degraded, many species employ cache protection strategies such as ceasing caching in the presence of others or avoiding storing perishable items for long periods. Several species frequently recover their caches and recache, which may reduce pilferage or degradation of cached items. We studied the food handling decisions of Florida scrub‐jays (Aphelocoma coerulescens) after cache recovery to determine the roles that social and ecological environments play in post‐recovery decisions. Instead of reducing recaching in the presence of others, recovering jays flew away from the recovery site, allowing them to eat or recache a recovered item regardless of the social context. Microhabitat type and soil moisture of the recovery sites had a significant influence on whether recoveries were eaten or recached; most items that were recached had been recovered from bare sand sites or sites with low soil moisture. Taken together, our results suggest that food store management of Florida scrub‐jays are unaffected by the social context, but are strongly affected by the habitat conditions that influence the quality of caches.     

贮食动物的盗食与反盗食行为策略

食物贮藏是许多动物应对食物短缺、保障其生存和繁衍的一种适应性行为。保护好贮藏食物以供食物短缺期利用,是食物贮藏成功的标志和进化动力。同种或异种动物盗食是贮藏食物损失的重要原因。嗅觉、视觉与空间记忆、随机搜寻等是动物搜寻和盗取食物的重要手段;避免盗食、阻止盗食和容忍盗食是动物反盗食的重要策略。动物通常采用多种行为策略进行盗食和反盗食,分配食物资源,形成相对稳定的种内、种间关系。盗食与反盗食互作及其对贮食行为进化的意义已成为行为生态学的研究热点和前沿之一,针对鸟类和哺乳类动物的研究尤为丰富。本文总结了贮食动物常见的盗食和反盗食行为策略及其相互作用的研究进展,主要内容涉及贮食动物利用嗅觉、视觉与空间记忆、随机搜寻等盗取其他个体食物的盗食策略,以及通过隐藏、转移、保卫、容忍等方式减少被盗食,保护贮藏食物的行为策略。针对现有研究状况,从种间盗食与反盗食及其与物种共存的关系,种间非对称盗食关系及其适应意义,盗食与反盗食最适行为策略及其与贮食动物适合度的关系等方面对今后研究提出了建议。     

Observational learning and the raiding of food caches in ravens, Corvus corax: is it ‘tactical’ deception?

Group-foraging ravens scatter-hoard when they are competing for food and, to some extent, also raid the caches made by others. We investigated the effects of observational spatial memory on individual caching and raiding tactics. With captive ravens, we found visual observation was essential for locating and raiding the caches of conspecifics. Both captive and free-ranging ravens, food cachers as well as potential cache raiders, responded to each other's presence. Cachers withdrew from conspecifics and most often placed their caches behind structures, obstructing the view of potential observers. Raiders watched inconspicuously and kept at a distance to cachers close to their cache sites. In response to the presence of potential raiders or because of their initial movements towards caches, the cachers frequently interrupted caching, changed cache sites, or recovered their food items. These results suggest that ravens, regardless of whether they act as cachers or raiders, are capable of withholding information about their intentions and, hence, manipulate the other bird's attention either to prevent or to achieve social-learning opportunities. Such interactions may qualify as ‘tactical’ deception and may have created a considerable pressure selecting for social cognition in ravens. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.