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Frugivory (animals ingest all or part of fleshy-fruits and defecate or regurgitate seeds) and scatter-hoarding (animals store seeds in small caches in soil) are two important vertebrate-mediated seed dispersal syndromes. In both instances, there is an exchange of food for seed transport, but there are many important differences. For example, the seeds of frugivore-dispersed plants are often clumped in feces or under roosts and the microsite of depostion is often unsuitable for seedling establishment. However, frugivores can transport seeds long distances allowing plants to colonize new areas and promoting gene flow between populations. Scatter-hoarding animals, on the other hand, actively space seeds in microsites that often favor seedling establishment, but don’t move seeds very far. Both modes of seed dispersal can be very effective, but in different way. Some species of plants have evolved propagules that take advantage of both modes of dispersal in two separate phases (i.e., diplochory), apparently to obtain both types of benefits.

Western chokecherry (Prunus virginiana var. demissa, Rosaceae) is dispersed by frugivorous birds and carnivores, but it has large seeds that are potentially attractive to rodents that could act as seed predators and dispersers. Here, we quantify the benefits of primary dispersal by birds and secondary dispersal by scatter-hoarding rodents. In the fall, avian frugivores (mostly American robins, Turdus migratorius, and cedar waxwings, Bombycilla cedrorum) consumed 87% of the fruit crop and dispersed 67% of the fruit crop away from parent plants. Rodents removed 89% of seeds that simulated bird-dispersed seed rain from transects in riparian zones and 58% from transects in upland habitats. Rodents scatter-hoarded 91.6% of the seeds they removed, burying most in small caches (two to eight seeds) 8-25 mm deep. About 39% of the seeds in spring caches produced seedlings. Inside rodent-proof exclosures, 52.1% of seeds buried to simulate rodent caches produced seedlings, 29.7% of which were still alive after 1 year. In contrast, only 3.8% of seeds placed on the soil surface, simulating dispersal by avian frugivores, produced seedlings. Seed dispersal by frugivorous birds likely contributes to colonization of unoccupied habitat through long-range dispersal and to escape from distance-dependent seed mortality near the parent plant. Despite seed losses, rodents offer short-range seed dispersal and bury seeds in more favorable sites for germination, improving seedling emergence and establishment. The combined mechanisms of seed dispersal significantly enhanced chokecherry seedling recruitment by providing more dispersal-related benefits than either frugivorous bird or scatter-hoarding rodents could provide alone.

Summary 1. The seeds of many plants are dispersed by animals, but the nature of these plant–animal mutualisms is often moulded by the abiotic environment. Here, we show that desert peach (Prunus andersonii), with dry fruits and large seeds, relies on scatter-hoarding rodents for dispersal and that this form of seed dispersal maintains the effectiveness of dispersal while reducing water expenditure. 2. The fruit pulp of desert peach dries and dehisces at maturity in early summer. No vertebrate frugivores consumed the fruits, but rodents quickly harvested the nuts. Nearly 75% of the nut crop was removed from plant canopies by scansorial rodents like white-tailed antelope squirrels (Ammospermophilus leucurus), which scatter-hoarded nuts in 1–2 seed caches 10–30 mm deep. Deer mice (Peromyscus maniculatus) also made many shallow one-nut caches. 3. Heteromyid rodents (Great Basin pocket mouse, Perognathus parvus; Panamint kangaroo rat, Dipodomys panamintinus) primarily larder-hoarded nuts (c. 60%) in burrows too deep for seedling emergence, but also scatter-hoarded nuts in a few, large caches 10–40 mm deep. 4. Antelope squirrels were the most effective dispersers and deer mice and Panamint kangaroo rats were less effective. Abiotically dispersed nuts had virtually no recruitment (

Food stored by animals can either be recovered by the hoarder or pilfered, but it is very difficult to distinguish between these possibilities under natural conditions. We tested the hypothesis that yellow pine chipmunks, Tamias amoenus, have a recovery advantage (i.e. a greater probability of finding a cached item than does a naive forager). We allowed chipmunks to cache radiolabelled pine seeds and then established artificial caches nearby. Paired artificial and real caches were identical in number of seeds, depth and microsite. Caches (393 pairs at 10 sites) were monitored during autumn 1997, 1999 and 2002. Removal of real caches represented the activity of the hoarder, foraging using spatial memory, plus the activities of pilferers, whereas the removal of artificial caches represented the activities of pilferers. We used these two measures in a competing risks model, which estimated that seed-caching chipmunks retrieved 74% of the food they stored in 1997 and 56% of the food they stored in 1999–2002. Seed-hoarding chipmunks were removed at four of the 10 sites immediately after they prepared caches. Where the seed-hoarders remained, animals took real caches 3.4 to 6.5 times faster than artificial caches, an effect that we attribute to the activity of the hoarder. Where the caching chipmunk was removed, there were no significant differences in the rates of removal of real versus artificial caches. We suggest that artificial caches can be used to estimate the proportion of caches retrieved by a seed-hoarding animal versus the proportion discovered by pilferers.

Many studies of postdispersal seed fate use seed removal as an index of seed predation. However, following primary seed dispersal, some seeds are transported intact by ants, dung beetles, scatter-hoarding animals, or abiotic processes to new microsites (secondary dispersal) where germination is possible. Despite a growing realization that secondary seed dispersal can play an important role in plant recruitment, many researchers continue to use seed removal as a proxy for seed predation and are focused too intently on only the initial step of seed fate. We describe, using examples from the recent literature, how the results of some seed removal studies may have been misinterpreted, present plausible, alternative explanations for the fate of seeds in those studies, and discuss the importance of detailed studies of seed fates. Following the fates of seeds can be difficult, but such studies contribute much more to our understanding of seed dynamics and plant fitness.

Olfaction plays an important role in the foraging behavior of rodents, but little is known about how this sense varies among taxa that evolved in different ecological settings. Field experiments were conducted in 10- by 10-m enclosures to test for interspecific differences in foraging success for ponderosa pine (Pinus ponderosa) or sunflower seeds, based on olfaction, of 4 rodent species: 2 heteromyid rodents (Panamint kangaroo rat, Dipodomys panamintinus, and Great Basin pocket mouse, Perognathus parvus) and 2 nonheteromyid rodents (yellow pine chipmunk, Tamias amoenus, and deer mouse, Peromyscus maniculatus). All species found large, shallow caches more frequently than small, deep caches. Chipmunks had the lowest foraging success for buried sunflower seeds under dry conditions. Deer mice did as well as the heteromyid rodents on large shallow caches but found relatively few small, deep caches. The heteromyids were the only species that found small, deep sunflower seed caches under dry conditions, and Panamint kangaroo rats always found more caches than did Great Basin pocket mice. These results confirm that olfaction plays an important role in foraging of rodents for buried seeds and support the notion that the olfactory sense of rodent species has evolved greater sensitivity to seeds in different ecological contexts.

We used a hierarchical approach to describe habitat characteristics of song posts and foraging sites used by Varied Thrushes (Ixoreus naevius) in coastal redwood (Sequoia sempervirens) forests of northwestern California. We measured mesohabitat (0.04-ha circular plots) and microhabitat (0.5-m radius) scale attributes centered on occupied and random song posts and foraging locations at four study sites during March–August of 1994 and 1995. Ninety-five percent of song posts were in trees or snags. Male thrushes used song posts with low foliage density near the top of large conifers (microhabitat scale), located on steeper slopes, surrounded by a high density of trees, and centered in drainages closer to water (mesohabitat scale) as compared to random locations. Varied Thrushes foraged predominantly on the ground early in the breeding season, then subsequently included fruit in their diet after the young had fledged. Although many variables were correlated with ground foraging locations, microhabitat foliage density had the greatest explanatory power, indicating thrushes selected foraging locations primarily at the microhabitat scale, and emphasizing the importance of measuring habitat characteristics at the appropriate spatial scale. Abrupt forest edges, such as those produced by clearcuts, may reduce habitat suitability for Varied Thrushes possibly explaining their absence from small forest fragments during the breeding season.