Shin Abe

SYNCHRONIZED ANNUAL SEED PRODUCTION BY 16 PRINCIPAL TREE SPECIES IN A TEMPERATE DECIDUOUS FOREST, JAPAN

To investigate synchronized annual fluctuation of seed production and its advantage for regeneration at the community level, for nine years (1987–1995) we monitored the flowering, seed production, and seedling emergence of the 16 principal tree species in a temperate deciduous forest, Ogawa Forest Reserve, in central Japan. We found that the species with higher synchronized flowering within a population had larger fluctuation of annual seed production at the population level. The coefficient of concordance of flowering and the coefficient of variation of annual seed production were continuously distributed among species, making it difficult to distinguish masting from nonmasting species. The annual seed production patterns of the 16 species were classified, by cluster analysis, into groups that synchronize their fluctuation of annual seed production. This analysis showed a highly synchronized annual seed production, not only among congeneric species, but also among species of different families. Although our results have some insufficiency of statistical significance, they did show that predator satiation, both in a population and a guild, effectively operated for many species to enhance seed survival at the pre-dispersal stage. They also showed that pollination efficiency was likely to be operating at the population level for half of the wind-pollinating species. However, generalist predator satiation at the postdispersal seed stage may not operate in a simple, detectable manner in this species-rich forest community. It is highly probable that there are combined effects of several factors: limited weather triggers for flowering, common flowering physiology among taxonomically related species, and the ecological advantages at the population and guild levels, may cause multiple species to have synchronized fluctuation patterns of seed production.

Community dynamics of Ogawa Forest Reserve, a species rich deciduous forest, central Japan

Forest community dynamics were studied for 4 years in a 6 ha permanent plot of species rich, old-growth, temperate deciduous forest in Ogawa Forest Reserve, central Japan. The gap formation rate, recruitment, mortality, gain and loss rate in basal area during 4 years were 42 m2 ha−1 yr−1, 1.74% yr−1, 1.19% yr−1, 1.12% yr−1 and 0.88% yr−1, respectively. The turnover time calculated from them ranged from 58 to 240 years. Both the mortality and mortality factors were size dependent; trees in middle size class had smallest mortality, and the proportion of the trees killed by disturbances increased with size. Gap creations were concentrated in a particular year, suggesting a large heterogeneity in time. Spatial distribution of recruited trees were biassed to the old gaps (older than 4 years), especially that of the species with Bell-shaped dbh distribution (shade intolerant) strongly associated with the gaps. Recruitment in tree stems and the loss of basal area, thus had the larger variability than mortality of stems and this forest, and the species with L-shaped dbh distribution seemed to going to increase the importance in the future if the present trend continues to be held. The turnover time of population is positively correlated with the maximum dbh size of the species, indicating the slow change of the population of large sized species.

Factors influencing sapling composition in canopy gaps of a temperate deciduous forest

To detect the factors that affect sapling species composition in gaps, we investigated 55 gaps in an old-growth temperate deciduous forest in Ogawa Forest Reserve, central Japan. Gap size, gap age, gap maker species, topographic location, adult tree composition around gaps, and saplings of tree species growing in the gaps were censused. For gaps ⩾5 m2, mean gap size was 70 m2 and the maximum was 330 m2. Estimated ages of gaps had a tendency to be concentrated in particular periods relating to strong wind records in the past. The sapling composition in gaps was highly and significantly correlated to that under closed canopy, indicating the importance of advance regeneration in this forest. However, some species showed significant occurrence biases in gaps or under closed canopy, suggesting differences in shade tolerance. The result of MANOVA showed that gap size and topography were important factors in determining the sapling composition in gaps. Species of gap makers affected the sapling composition indirectly by influencing gap size. The existence of parent trees around gaps had effects on sapling densities of several species. Gap age did not have clear influences on sapling composition. Variations in gap size and topography were considered as important factors that contribute to maintenance of species diversity in this forest.

Effect of Mast Production on Home Range use of Japanese Black Bears

ABSTRACT Although bears may expand their home ranges in times of low food availability, it is unclear what mechanisms directly affect home range extension in times of low mast production in Japanese forests. To clarify the relationship between home range utilization by Japanese black bears (Ursus thibetanus) and abundance and distribution of mast production, we collected data on habitat use from 13 bears (6 M and 7 F) fitted with Global Positioning System (GPS) collars equipped with activity sensors in the Ashio-Nikko Mountains on the eastern part of Honshu Island, Japan, during 2006–2008. We also collected data on mast production by 5 Fagaceae species. We categorized each fall as either poor (2006) or relatively-good (2007 and 2008) based on mast production. Bears used small patches in their large home ranges and the distances between core areas increased in the fall of 2006, when the mast of Japanese oak (Quercus crispula) were poorly distributed. Our findings suggest that localized patches of Japanese oak are the staple food for bears in our study area, even in poor mast years. However, in the fall of 2006, we also found that bears moved to lower elevations, relative to 2007 and 2008, in search of alternative foods (e.g., Konara oak [Q. serrata] and Japanese chestnut [Castanea crenata]), which were mostly at lower elevations and produced mast consistently over the study period. Our results suggest that dispersion and elevational distribution of mast-producing trees affect bear habitat use in fall, as well as amount of mast.

Effects of canopy gaps on the demography of the subcanopy tree Styrax obassia

To evaluate the gap dependency of the subcanopy tree species Styrax obassia, we estimated the demographic parameters of this species in a permanent plot in the Ogawa Forest Reserve, Central Japan, a temperate deciduous forest. From the data collected over eight years, we constructed four transition matrix models of Styrax: a whole population; a shaded subpopulation; a gap-site subpopulation; a compound population of shaded and gap-site subpopulations. The whole- population model suggested that the population structure of Styrax is stable in its present condition. The asymptotic popu- lation growth rate of the shaded subpopulation was larger than 1, suggesting that this species can maintain its population even under shaded conditions. The elasticity analyses showed the relative importance of survival of adult individuals that stay mostly under a closed canopy. Since the subpopulation in gaps is very small in a mature forest, the contributions of high seedling survival and high fecundity of adult trees in gaps to the whole population growth were smaller than the survival of adult individuals in the shade. Although we showed that this species has the potential to increase its population in larger disturbances, it actually dominates in the study site where the disturbance scale is relatively small. Styrax should be classi- fied as a typical shade-tolerant species, and it also takes good advantage of small scale disturbance. It can sufficiently main- tain its population in a mature forest with small-scale distur- bances and has the adaptive characteristics of species that live under a closed canopy.

Comparative demography of three coexisting Acer species in gaps and under closed canopy

Abstract Questions: 1. Is there a trade-off between gap dependency and shade tolerance in each of the life-history stages of three closely related, coexisting species, Acer amoenum (Aa), A. mono (Am) and A. rufinerve (Ar)? 2. If not, what differences in life-history traits contribute to the coexistence of these non-pioneer species? Location: Ogawa Forest Reserve, a remnant (98 ha), species-rich, temperate deciduous forest in central Japan (36°56′ N, 140°35′ E, 600 - 660 m a.s.l.). Methods: We estimated the demographic parameters (survival, growth rate and fecundity) by stage of each species growing in gaps and under closed canopy through observations of a 6-ha permanent plot over 12 years. Population dynamics were analysed with stage-based matrix models including gap dynamics. Results: All of the species showed high seedling and sapling survival rates under closed canopies. However, demographic parameters for each growth stage in gaps and under closed canopies revealed inter-specific differences and ontogenetic shifts. The trade-off between survival in the shade and growth in gaps was detected only at the small sapling stage (height < 30 cm), and Ar had the highest growth rate both in the shade and in the gaps at most life stages. Conclusions: Inter-specific differences and ontogenetic shifts in light requirements with life-form differences may contribute to the coexistence of the Acer species in old-growth forests, with Aa considered a long-lived sub-canopy tree, Am a long-lived canopy tree, and Ar a short-lived, ‘gap-phase’ sub-canopy tree. Nomenclature: Satake et al. (1989).

Effects of abiotic and biotic factors and stochasticity on tree regeneration in a temperate forest community.

Abstract: How deterministic or stochastic is tree regeneration in a temperate forest? To answer this question, we analyzed the relative effects of abiotic and biotic factors on seed and seedling demography of tree species in an old-growth temperate forest, Ogawa Forest Reserve, Japan. The working hypothesis was that seed and seedling demography of trees would be controlled more deterministically in comparison with tropical forests and that the magnitude of deterministic effects of the abiotic and biotic factors would depend on regeneration traits relating to seed dispersal ability. We examined 5 abiotic factors (litter and humus cover, vegetation cover, light level, soil moisture, and relative elevation) and 2 species-specific biotic factors (distance from the nearest conspecific adult and density of conspecific seeds or seedlings) to quantify their contribution to the spatial variation in seedling emergence and survival for 18 principal tree species in the community. The results showed that conspecific seed density had a negative effect on seedling emergence for almost all species, as suggested by the Janzen—Connell model. On the other hand, various factors were detected for seedling survival of each species. We also found that regeneration traits relating to seed dispersal ability were significantly correlated with some of the abiotic factors. However, the 7 factors tested were less important than expected; seed and seedling demography appeared to be determined rather stochastically in this stable forest community. Even in a temperate forest, seed and seedling demography appeared to be rather stochastic, and niche partitioning among species during these stages was of limited importance.

Effect of hard mast production on foraging and sex-specific behavior of the Asiatic black bear (Ursus thibetanus)

Abstract. We studied the relationships between movement and foraging habits of the Asiatic black bear (Ursus thibetanus) and hard mast production of five tree species in cool temperate forest during 2006–2008. The purpose of this study was to test the hypothesis that low mast production affects sexual differences in bear behavior. We summarized the movement pattern of 13 bears in terms of minimum movement distance and cumulative movement distance of the movement path followed during 24 hour. Masting of Quercus crispula acorns was low in 2006, high in 2007 and moderate in 2008. The dominant food items found in bear scats were hard mast, especially Quercus acorns. The percentage of Quercus acorns in the food items in scats was higher in 2007 than in 2006 and 2008. Movement distance of males and females increased in the low mast year. However, the increase of movement distance of females was larger than that of males. Thus, masting influenced the behavior of females more strongly than males. Our results indicated that low mast production changed the food habits and the size of the home range of bears, especially of females.

Relationship between the decomposition process of coarse woody debris and fungal community structure as detected by high-throughput sequencing in a deciduous broad-leaved forest in Japan.

We examined the relationship between the community structure of wood-decaying fungi, detected by high-throughput sequencing, and the decomposition rate using 13 years of data from a forest dynamics plot. For molecular analysis and wood density measurements, drill dust samples were collected from logs and stumps of Fagus and Quercus in the plot. Regression using a negative exponential model between wood density and time since death revealed that the decomposition rate of Fagus was greater than that of Quercus. The residual between the expected value obtained from the regression curve and the observed wood density was used as a decomposition rate index. Principal component analysis showed that the fungal community compositions of both Fagus and Quercus changed with time since death. Principal component analysis axis scores were used as an index of fungal community composition. A structural equation model for each wood genus was used to assess the effect of fungal community structure traits on the decomposition rate and how the fungal community structure was determined by the traits of coarse woody debris. Results of the structural equation model suggested that the decomposition rate of Fagus was affected by two fungal community composition components: one that was affected by time since death and another that was not affected by the traits of coarse woody debris. In contrast, the decomposition rate of Quercus was not affected by coarse woody debris traits or fungal community structure. These findings suggest that, in the case of Fagus coarse woody debris, the fungal community structure is related to the decomposition process of its host substrate. Because fungal community structure is affected partly by the decay stage and wood density of its substrate, these factors influence each other. Further research on interactive effects is needed to improve our understanding of the relationship between fungal community structure and the woody debris decomposition process.

Fluctuation of daily activity time budgets of Japanese black bears: relationship to sex, reproductive status, and hard-mast availability

Abstract We documented the fluctuation of daily time budgets in Japanese black bears (Ursus thibetanus japonicus) throughout the year using continuous day-to-day data from activity sensors integrated into global positioning system collars, during 2003–2009. We evaluated the influence of availability of food resources on daily active time of bears in conjunction with differences in sex and reproductive status of females (with or without offspring). The daily active time of bears fluctuated nonlinearly throughout the year. There were 3 turning points at which the activity level clearly changed from increasing to decreasing, or vice versa. Level of activity gradually increased in spring and reached a peak in July, then decreased and reached a trough in late August, and increased and reached a peak again in October. Males and females exhibited similar patterns of seasonal fluctuation in daily active time, although there were differences of activity levels between sexes during some periods. Dates of the 3 turning points did not differ between sexes, or among years. Seasonal variation in food availability may explain the timings of the turning points, at least in part. In addition, yearly variation in food resources, especially hard mast, may have affected the increasing and decreasing pattern of daily active time during autumn. Our results suggest that evaluating activity level based on pooled data without examining differences within a season (or month), and differences in year, sex, and reproductive status may result in misinterpretation.