Teruaki Hino

How do deer affect tree seedlings on a dwarf bamboo-dominated forest floor?

A field experiment was conducted in Mt. Ôdaigahara, west-central Japan, to reveal the effects of two herbivores, Sika deer (Cervus nippon) and mice (Apodemus spp.), on the seedlings of five cohorts of three tree species, Abies homolepis, Fraxinus lanuginosa f. serrata and Fagus crenata. The forest floor of the study site was covered with dwarf bamboo, Sasa nipponica, which would also affect the seedlings. Eight combinations of three treatments were set: exclusion of deer, exclusion of mice and removal of dwarf bamboo. Deer were expected to affect the seedlings not only negatively by foraging but also positively by browsing dwarf bamboo that overshadows seedlings. The survival of these cohorts was analyzed by survival analysis and the differences in their survival between the treatments were investigated. The results of the experiment showed that: (1) exclusion of deer increased the aboveground biomass of dwarf bamboo and made it more difficult for seedlings to survive under the regenerated dwarf bamboo stand, and (2) deer negatively affected the seedlings, but they had positive, indirect effects in some cohorts by decreasing the aboveground biomass of dwarf bamboo. However, such effects were not detected in some cohorts which were affected excessively by deer or dwarf bamboo. No clear results on the effect of mice on seedlings were obtained. We conclude that a positive, indirect effect of deer on seedling survivorship would be observable when the magnitude of a positive, indirect effect caused by decreasing dwarf bamboo is larger than that of a negative, direct effect of deer.

Effects of deer, mice and dwarf bamboo on the emergence, survival and growth of Abies homolepis (Piceaceae) seedlings

In the subalpine mixed forest of Mt. Ôdaigahara, mid-western Japan, the understory is dominated by dwarf bamboo (Sasa nipponica), which is the major forage of overly populous sika deer (Cervus nippon). In the present study, we monitored the survival and growth of Abies homolepis seedlings over 5 years to determine how they responded to the experimental removal of dwarf bamboo and to the exclusion of sika deer and mice (Apodemus argenteus and A. speciosus). Deer and dwarf bamboo reduced the survival of seedlings but had different effects on growth. The stems of seedlings were shorter in the presence of deer, indicating that taller seedlings were apt to be browsed by deer, whereas the diameters of seedlings were smaller in the presence of dwarf bamboo, mainly owing to its shading effect. The presence of mice decreased the number of seedlings germinating in a particular site, but had no effect on seedling survival after germination. There was no significant indirect effect whereby the survival of seedlings was predicted to be facilitated by the decreased biomass of bamboo because of grazing by deer. We supposed that this might be because the direct negative effect of deer was so large as to conceal the positive indirect effect.

Dwarf bamboo as an ecological filter for forest regeneration

Quantitative relationships between the aboveground biomass of dwarf bamboo, Sasa nipponica, and the survivorship and emergence of seedlings of Abies homolepis, Fraxinus lanuginosa f. serrata, and Fagus crenata were estimated. We show that dwarf bamboo acts as an ecological filter since the responses of tree species differ according to the biomass of dwarf bamboo. Deer exclusion without management of dwarf bamboo would make it impossible for any tree species to regenerate due to rapid increases in the biomass of dwarf bamboo.

Prey distribution, foliage structure, and foraging behavior of insectivorous birds in two oak species (Quercus serrata and Q. variabilis)

We examined foraging behaviors of four insectivorous bird species (Great Tit Parus major, Varied Tit P. varius, Long-tailed Tit Aegithalos caudatus and Japanese White-eye Zosterops japonica) on the two oak species (Q. serrata and Q. variabilis) in relation to fine-scale foliage structures, abundance, and distribution of folivorous insects and physical and chemical characteristics of leaves. There was no difference in the mean number of insects for each oak species, but there was a significant difference in the mean biomass. The distance from twigs or branches to leaf tips or blade-petiole junctions showed that leaves of Q. variabilis were more dispersed than those of Q. serrata. The different size distribution of insects influenced insect sizes selected and foraging maneuvers used by each bird species. Parus major and P.varius fed on beetles (low energy gain) with perch-gleaning (the most energy-saving maneuver) from the upper side of leaves on Q. variabilis, whereas they fed on large lepidopteran larvae (high energy gain) from the lower side of leaves on Q. serrata using a variety of maneuvers (including energy-consuming ones). Both A. caudatus and Z. japonica fed on insects from the lower side of leaves by hang-gleaning and reach-gleaning, respectively. These species appeared to be morphologically adapted to the energy-consuming but more effective maneuvers used to take preys from the undersides of leaves. We hypothesize that differences in foliage structures between closely related species of trees have evolved as a balance between defense against folivorous insects and attraction of insectivorous birds.

Deer Browsing on Dwarf Bamboo Affects the Interspecies Relationships among the Parasitoids Associated with a Gall Midge

We found that deer browsing alters the species composition of parasitoids of gall-forming insects via its effects on the host plants. At Mt. Odaigahara, in west-central Japan, we compared the species composition of two parasitoid wasps, Pediobius sp. (Eulophidae) and Torymus sp. (Torymidae), on an unidentified gall midge (tribe Oligotrophini) that forms galls on dwarf bamboo (Sasa nipponica Makino et Shibata), the major forage for Sika deer (Cervus nippon Temminck). Gall width was larger inside deer exclosures, where the bamboo culms were longer and thicker due to their escape from browsing. The parasitism rate by Pediobius sp. was lower inside the exclosures, where parasitism concentrated on the smaller galls. In contrast, the parasitism rate by Torymus sp. was higher inside the exclosures and concentrated on the larger galls. Torymus sp. emerge earlier and have a longer ovipositor than Pediobius sp., thus it should be able to oviposit throughout all gall developmental stages. Because Torymus sp. may be hyperparasitized by Pediobius sp., more Torymus sp. larvae survived in larger galls that Pediobius sp. could not penetrate with its shorter ovipositor. Thus, deer browsing indirectly favors Pediobius sp. by reducing gall size and thereby improving access to host.

Nitrogen leaching from surface soil in a temperate mixed forest subject to intensive deer grazing

We aimed to determine the grazing impact on soil nitrogen (N) mineralization and N leaching in a temperate mixed forest on Mt. Ohdaigahara, central Japan, where Sika deer (Cervus nippon Temminck; hereafter deer) grazing has decreased biomass of Sasa nipponica Makino et Shibata (hereafter sasa) that dominates the forest understory. Soil N mineralization and N leaching of the top 0.3-m layer were measured in situ from 2006 to 2007 by an ion-exchange resin core method inside and outside a deer exclosure. Deer grazing decreased the carbon to nitrogen (C:N) ratio of sasa litter because of the declining dead culm C:N ratio and the increasing ratio of leaves with a lower C:N ratio than culms in the aboveground biomass. Grazing increased N mineralization and nitrification rates as well as annual N (ammonium-N and nitrate-N) and nitrate-N leaching. In contrast, deer grazing did not increase the total soil C and N content. The positive effect of grazing on the N mineralization rate may have resulted from a qualitative change in sasa litter due to grazing. The higher annual N leaching resulting from deer grazing may have been related to the facilitated N mineralization rate and decreased N uptake in plants, e.g., sasa with less aboveground biomass due to grazing.