Five members of the order Carnivora in the cool-temperate deciduous forests of Japan consume the fruits of trees, and they are potential dispersers of the seeds of fleshy-fruited plants. We studied the frugivory of the Asiatic black bear, Japanese marten, badger, red fox, and raccoon dog in cool-temperate deciduous forest of central Japan. From May 2003 to April 2005, a total of 377 fecal samples of the five carnivores (bears, 91; martens, 158; badgers, 45; foxes, 36; and raccoon dogs, 47) were sampled to determine the presence and frequency of occurrence of seeds. Seeds from 17 plant species, representing about 50% of the fleshy-fruited plants occurring in the study forest, were recovered from the carnivore fecal samples. Large numbers (9-10,256) of seeds were present in those feces. Almost all the seeds of fleshy fruits retrieved from the fecal samples were undamaged whereas no intact acorns or nuts were recovered. These findings suggest that all five carnivores can act as seed dispersers for some fleshy-fruited plants in cool-temperate deciduous forest.
We estimated the seed shadow created by the Asiatic black bear Ursus thibetanus in order to evaluate the bears eff ectiveness as a seed disperser. We combined data from bear movements, determined by GPS telemetry, with data from gut retention time (GRT). We estimated plant seed shadows in two ways: from direct movement data to give the actual seed shadow (ASS), and from cumulative movement data to give the potential seed shadow (PSS). Th e purpose of this study was to answer the following questions: (1) does GRT diff er between seasons or in the size of contents? (2) Does seed shadow vary among sex, seasons, estimation method (ASS or PSS), and years? (3) Does the masting aff ect seed shadows? Th ere were no diff erences in median GRT among seasons or seed dimensions. Combining these data, the seed shadows produced by long GRT (median; 15.2 -19.7 hours, maximum; 44.0 hours) and large daily movements suggest that the bears eff ectively move 40% of the seeds they consume to a distance greater than 500 m from the parent tree and can potentially move the seeds up to a maximum distance of more than 22 000 m from the parent tree. Th e results also indicate that bears make complex seed shadows caused by multiple defecations and long periods of daily movement. In summary, PSS did not diff er between sexes, but PSS can be expected to be larger in autumn than in summer of each year. ASS, however, can be expected to be larger in males than females, and to be larger in autumn than in summer. ASS may become especially large during a poor masting year as compared to good masting years. Th ese results indicate that bears are potentially more eff ective seed dispersers during years of poor fruit production in autumn. Th e bears have longer seed shadows than other seed disperser and consequently may play a unique role in the maintenance and renewal of forest ecosystem.
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