Effects of food value, predation risk, and pilferage on the caching decisions of Dipodomys merriami

Animals that scatter cache their food face a trade-off betweenthe benefits of protecting caches from pilferers and the costsassociated with caching. Placing food into a large number ofwidely spaced caches helps to protect it from pilferage butalso involves costs such as greater exposure to predators. Ipredicted that animals would disperse food into a larger numberof more widely spaced caches when caching (1) a preferred foodversus a less preferred food and (2) under conditions of lowpredation risk versus high predation risk. To test these predictions,I examined the scatter-caching decisions of Merriam's kangaroorats (Dipodomys merriami). D. merriami distributed caches inclumped patterns, regardless of food preference, but they showeda tendency to invest more in a preferred food by distributingcaches more widely. Under the relative safety of the new moon,they did not disperse caches more widely, rather they partitionedthe same amount of food into a larger number of caches thanthey did under the full moon, when predation risk is higher.To examine whether their cache spacing decisions had a significantimpact on the success of cache pilferers, I measured discoveryby pilferers of artificial caches of two food types at differentcaching distances. Results indicate that the cache spacing behaviorof D. merriami functions to protect caches from pilferers, becauseincreased spacing of artificial caches decreased the probabilityof pilferage for both types of food.     

Spatial patterns of food storage by Merriam's kangaroo rats

We investigated scatter-hoarding behavior by Merriam's kangaroorats (Dipodomys merriami) in a complex arena in the laboratory.Animals cached seeds in four sand-filled boxes connected byrunways, with an artificial burrow at one end of the systemand a food source at the other. Animals initially cached moreseeds in boxes close to the food source, but amounts cachedbecame more evenly distributed among boxes as sessions progressed.The results were consistent with the hypothesis that scatterhoarding by kangaroo rats may represent a compromise betweenrapidly sequestering seeds, hence making them unavailable tonondigging competitors, and spacing-out caches, making themless vulnerable to pilferage by other 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.     

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.     

Evidence of Social Influences on Cache‐Making by Grey Squirrels (Sciurus carolinensis)

Many studies have found that scatter‐hoarding animals change their behaviour when storing food in the presence of conspecifics to minimize the likelihood that their caches will be pilfered; they refrain from caching, move away from conspecifics or choose visually obscured sites. This study reports the first evidence that the presence of conspecifics continues to influence the caching behaviour of a scatter‐hoarding mammal, the grey squirrel, after a suitable cache site has been selected and the hoarder is filling and covering its cache. Wild grey squirrels were filmed when storing preferred and less preferred nuts and when they were alone or with conspecifics present. In line with previous findings, squirrels spent longer travelling from the nut patch and were more vigilant when conspecifics were present. However, squirrels also spent longer disguising their caches and were more likely to stop digging and become vigilant when conspecifics were present than when they were alone. In particular, they were most likely to curtail their digging when storing their preferred nuts in the presence of conspecifics. The results indicate that caching squirrels remain sensitive to the presence of conspecifics until the cache is complete and that they respond flexibly to conspecifics according to the type of food they are storing.     

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.     

Reciprocal pilferage and the evolution of food-hoarding behavior

Current theories of food-hoarding behavior maintain that hoardingcan be adaptive if a hoarder is more likely than any other animalto retrieve its own caches. A survey of the literature indicatedthat the hoarder often has a recovery advantage when searchingfor items it has stored, but levels of cache pilferage are oftenso high (2–30% per day) that at least for some long-termfood hoarders, the caching animal is unlikely to recover a significantamount of its stored food. Except in a few cases (acorn woodpeckersand beavers), kin selection cannot explain the high levels ofpilferage observed. We suggest that some small solitary animalswith overlapping home ranges (e.g., most rodents, chickadees,and tits) are able to tolerate high levels of cache pilferage.Pilferage is not as damaging to these animals as it might otherwisebe because many interspecific and all intraspecific cache pilferersalso cache food. These or similar food caches can be pilferedlater by the original food hoarder. In other words, pilferingin these species is often reciprocal, and because it is reciprocal,it can be tolerated. We argue that caching systems based onreciprocal pilfering can be stable and are not necessarily susceptibleto "cheaters," animals that pilfer food but do not scatter hoardfood themselves, and we introduce a model of food hoarding tosupport this argument. These food-caching systems based on reciprocalpilfering resemble cooperative behavior, but the behavior isactually driven by the selfish interests of individuals. Thistheory of scatter-hoarding behavior based on reciprocity hasimportant implications for the ways that food-hoarding animalsinteract with inter- and intraspecific competitors.     

基于贮食行为的鸟类社会认知研究进展

贮食是动物应对环境变化和不可预测性而进化出的有效生存对策,认知则是当前鸟类学研究的热点问题之一。目前鸟类贮食行为中的认知研究多集中在空间认知,而社会认知研究相对滞后。对于贮食物种而言,储藏食物被盗现象非常普遍,为了避免被盗食,贮食者不仅要有发达的空间认知能力去记忆贮食地点,同时还需要极强的社会认知能力处理与盗食者的关系,可见社会认知在鸟类的贮食行为中扮演着重要角色。本文将从鸟类贮食的社会关系认知以及社会地位认知两个方面,对鸟类贮食行为中的社会认知研究进行综述,以期为后续鸟类社会认知研究提供借鉴和参考。     

啮齿动物对植物种子的多次贮藏

啮齿动物对种子的多次搬运和贮藏是极为复杂的行为过程,既是对自己贮藏物进行管理、防御竞争者盗食的一种策略,也是盗食其它个体贮藏物的一种食物利用方式。此外,啮齿动物多次贮藏种子的过程实际上也是植物种子的多阶段扩散过程,因而对植物更新产生重要影响。本文综述了啮齿动物对植物种子多次贮藏的研究进展,分析了多次贮藏种子的原因,并从啮齿动物与植物的相互关系上探讨其生态学意义。     

Seasonal variation in the effect of cache pilferage on cache and body mass regulation in Carolina chickadees: what are the trade-offs?

Current dynamic optimization models predict that animals shouldrespond to cache pilferage by decreasing the probability ofcaching food and by increasing internal fat storage to compensatefor a reduction in cache size. We tested these predictions underlaboratory conditions with variable food access (four 15-minintervals/day). Carolina chickadees (Poecile carolinensis) weresubjected to two environments: under pilferage conditions, one-quarterof their cached seeds were stolen every 0.5 h, and under no-pilferageconditions, seeds were left in place. Half the birds startedwith pilferage conditions and were then switched to the no-pilferagecondition; the other half started with no pilferage and werethen switched to pilferage conditions. The experiment was conductedover the course of a year to test for seasonal variation inthe response to seed pilferage. The birds responded to seedpilferage by taking more seeds from a feeder, suggesting thatthey monitored cache availability. Alternatively, the birdsmay have taken additional seeds from the feeder in responseto increased hunger caused by a loss of cached food. Contraryto our prediction, birds cached a higher percentage of seedsfrom the feeder when cached seeds were pilfered than when cacheswere left in place. Treatment order also affected caching behaviorfor all but the summer birds: chickadees initially subjectedto pilferage stored a higher proportion of seeds than thoseinitially subjected to no pilferage. Caching percentages inthe summer were unaffected by cache pilferage. Caching rates(number cached/day) also followed the same trends: rates werehigher when seeds were pilfered than when seeds were not pilfered,and there was a treatment-order effect for all but the summerbirds. Variation in body mass also failed to match predictedtrends. All birds exhibited a monotonic increase in mass asthe experiments proceeded, irrespective of treatment order.Controlling for this monotonic increase in mass, an analysisof residual variation in body mass indicated that birds gainedless weight when seeds were pilfered than when seeds were leftin place. Finally, birds tested in the fall and spring wereheavier than those tested in the summer. These results failto support the relationship between cache maintenance and bodymass regulation predicted by current models of energy regulation.We discuss the applicability of three hypotheses for the observedtrends.