The role of birds and mammals in Korean pine (Pinus koraiensis) regeneration dynamics.

This study examined the interrelationships of the fall seed-foraging guild with Korean pine (Pinus koraiensis) and its regeneration. The study took place in old-growth and secondary-growth forests in Northeastern China. Foraging behavior, seed transportation distance, and cache location of various birds and mammals were observed. Regeneration characteristics of Korean pine were also analyzed using plots in various vegetation cover types, successional stages, and topographical situations. Ten species of vertebrates were able to harvest seed from the closed-cone Korean pine. Of these, the Eurasian nutcracker, Eurasian nuthatch, red squirrel, and Siberian chipmunk were found to be potential seed dispersal agents. The nutcracker was the most important dispersal agent, easily acquiring seed with its large pointed bill, carrying up to 62 seeds in one trip, carrying seed at least 4 km, and placing seed in a variety of sites 2.5-3 cm deep in the soil. The Eurasian nuthatch carried a single seed per trip at distances less than 50 m, needed nutcrackers to open the cones and expose seed before they could acquire seed, and occasionally cached seed in the soil. Red squirrels were uncommon visitors to the tree tops of Korean pine, carried cones shorter distances than nutcrackers, and were only found under forest canopies. Human harvest of cones by knocking off branches also affected squirrel behavior and reduced future cone crops. Siberian chipmunks also collected seed from cones in trees and appeared to transport seed less than 50 m. Six other species were observed in this study successfully harvesting seed from cones but were not potential seed dispersers. Natural seedling establishment was found to be over 1000 seedlings/ha except on old-growth pine-hardwood sites. Squirrels were commonest here, but few seedlings survived past the 2nd year due to the intense shading. Second-growth forest types, including aPicea plantation where nutcrackers cached seed daily, and an old-growth pine-hardwood selective-harvest site, had the greatest regeneration. In conclusion, most natural regeneration of Korean pine in this part of its range is due primarily to the Eurasian nutcracker. Nutcrackers can aid forest managers in reaching desired stocking levels after disturbance, as well as a more natural-appearing forest. Squirrels, chipmunks, and nuthatches are minor seedling establishment agents. Korean pine seed is an important food source used by at least 22 species of forest wildlife.