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.     

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.     

The placement, recovery, and loss of scatter hoards by eastern chipmunks,Tamias striatus

Although eastern chipmunks typically larder hoard food in theirburrows, some scatter hoarding occurs, especially by newly emergedjuveniles and by females with young. To examine patterns ofscatter hoard placement and their adaptive significance, weused standard food provisioning tests followed by continuousmonitoring to determine the longevity and fate of scatter hoards.Longevity of scatter hoards was low (median 74 min) and pilferagewas high (46%). Chipmunks placed scatter hoards away from thepatch, closer to their burrow and within the defended area ofprimary use. Scatter hoard placement was affected by the numberof competitors at the patch. However, juveniles and femaleswith young differed slightly in their responses. competitorsthat approached scatteihoards were sometimes chased away bythe scatter hoard's owner, and the scatter hoard was moved toanother location by the owner. Females with young recoveredtheir largest scatter hoards first, but there was little additionalpattern in scatter hoard recovery. An experiment using simulatedscatter hoards showed that scatter hoards farther from a patchof food lasted longer and that disturbing and marking scatterhoards reduced their longevity. These observations indicate:(1) that scatter hoard placement is more related to avoidingpilferage of completed caches than to rapidly sequestering fooditems from an ephemeral patch and (2) that optimal cache distributionmay be affected by the cache-defense ability of individuals,by the relationship between competitor density and cache securitynear the patch, and by the territorial behavior of neighboringconspecifics.     

Regulation of cache stores and body mass in Carolina chickadees (Panes carolinensis)

Here I report the results of laboratory experiments on cachingbehavior of Carolina chickadees (Parus carolinensis) designedto test the following predictions from a recent dynamic optimizationmodel: under limited resources, small birds, when fat, shouldcache food instead of eating it, but they should eat insteadof caching when lean; when resources are abundant, birds shouldcache less when fat and more when lean. In addition, when resourcesare abundant the amount of time spent in foraging-related activityshould decrease with an increase in body mass. Chickadees weretested for 2 weeks in either a poor-quality environment (wherethey were given four 5-min periods per day of access to a feederfilled with sunflower seeds) or a richer environment (wherethey were given four 10-min access periods); they were thenswitched to the alternative environment for an additional 2weeks. The entire experiment lasted from October through June.Within-individual comparisons showed that birds in both thepoor and rich environments exhibited the predicted correlationsbetween the probability that a seed was cached and body massmeasured at dawn. The number of seeds already stored had a weakand variable effect on caching decisions. Time budgets changedas predicted; birds spent less time with food-related behaviorswhen their energy stores were high compared to when they werelow, and overall spent a higher fraction of their time foragingin the poor-quality environment compared to the rich environment.Two other variables also affected caching behavior: experienceand season. Individuals were more likely to cache in the poorenvironment when they had been tested in the rich environmentthe previous 2 weeks, as compared with birds started in thepoor environment. However, this was true only for tests donein the winter. In late spring, all birds stopped caching orreduced caching rates when faced with limited food availability,irrespective of previous experience. The birds harvested seedsfaster when they cached instead of eating seeds; as a result,long-term weight regulation in the poor environment was affectedby caching decisions. Birds tested in late spring lost weightin the poor environment, whereas those tested in the wintermaintained a stable weight when switched from the rich to thepoor environment. Thus, annual endogenous cues directly affectcaching decisions and indirectly influence the long-term regulationof body mass     

Testing the quick meal hypothesis: The effect of pulp on hoarding and seed predation of Hymenaea courbaril by red‐rumped agoutis (Dasyprocta leporina)

Abstract: Red‐rumped agoutis (Dasyprocta leporina) are important seed dispersers/predators of Neotropical large‐seeded plants. Several species of seeds cached by agoutis have an edible reward, in contrast to temperate rodent‐dispersed diaspores. The quick meal hypothesis states that the presence of a reward such as edible pulp will enhance the efficiency of rodents as seed disperses by satiating the animal and, consequently, reducing seed predation and enhancing hoarding. In this study, this hypothesis was tested using as the reference system the pulp and seeds of Hymenaea courbaril. Seeds with and without pulp were offered to agoutis and the behaviour of each individual was recorded. Since the probability of predation and hoarding were complementary, we used the probability of predation. The proportion of agoutis that preyed on at least one seed was similar for seeds with (42.8% of individuals) and without (40.0% of individuals) pulp. In agoutis that preyed upon at least one seed, the probability that they killed a seed did not differ between seeds with (0.17 ± 0.03) and without (0.20 ± 0.08) pulp. Hence, these results do not support the ‘quick meal hypothesis’.     

Cache site selection by chipmunks (Tamias spp.) and its influence on the effectiveness of seed dispersal in Jeffrey pine (Pinus jeffreyi)

The effectiveness of Jeffrey pine (Pinus jeffreyi) seed dispersal performed by seed-caching yellow pine chipmunks (Tamias amoenus) and lodgepole chipmunks (Tamias speciosus) was compared to that of wind dispersal in the Sierra Nevada of western Nevada. Wind-dispersed seeds typically fall under or near the parent tree. Chipmunks removed 90 and 97% of 1064 radioactive seeds from each of two simulated wind-dispersed seed shadows in less than 24 h. Wind-dispersed seeds were deployed within 12 m of the two source trees, but chipmunk caches were found from 2–69 m from the trees. Chipmunks carried nearly all seeds away from source trees, greatly reducing the density of seeds under and near source trees. Caches contained from 1–35 seeds and most were buried 7–21 mm deep. Chipmunks cached in open bitterbrush shrubland with mineral soils much more than expected and cached in closed-canopy Jeffrey pine and lodgepole pine forests with thick needle litter much less than expected. Many Jeffrey pine seedlings and saplings grow in the bitterbrush habitat and few grow in the pine forests. Ten and 20% of the original caches survived until April, the time of seed germination, at the two sites. The movement of wind-dispersed seeds is random relative to environmental variables important in seedling survival, and the wind in coniferous forests cannot quickly bury seeds. The quality of seed dispersal rendered by chipmunks was superior to that provided by the wind because the chipmunks quickly harvested seeds on the ground, moved them away from source trees, and buried them in the ground in habitats and microhabitats where they were more likely to establish new seedlings. The increased quality of seed dispersal provided by animals relative to the wind may help explain why over twenty species of pines have evolved seeds and cones that are adapted for dispersal by seed-caching animals.