Comparative study of leaf carbon gain in saplings ofThujopsis dolabrata var.hondai andQuercus mongolica var.grosseserrata in a cool-temperate deciduous forest

Seasonal courses of leaf CO₂ gas exchange in a growing season were examined in saplings ofThujopsis dolabrata var.hondai andQuercus mongolica var.grosseserrata in a cool temperate deciduous forest. Between the two tree species there were no large differences in the light compensation point of leaf photosynthesis, except for the season of new leaf expansion. However, light-saturated rates of net photosynthesis were obviously high inT. dolabrata var.hondai. EvergreenT. dolabrata var.hondai saplings had large photosynthetic production in two seasons, before the emergence of new foliage and after foliage fall of the overstory deciduous trees, because of the significantly high solar radiant energy penetrating under the forest canopy during the seasons. Saplings of deciduousQ. mongolica var.grosseserrata were heavily shaded throughout the growing season by foliage of the overstory trees, which resulted in a low daily surplus production. The annual surplus production of leaves in the growing season was estimated to be 2300 mmol CO₂ m⁻² inT. dolabrata var.hondai and −100 mmol CO₂ m⁻², slightly negative, inQ. mongolica var.grosseserrata. These results supported the high survivability ofT. dolabrata var.hondai saplings and the high mortality ofQ. mongolica var.grosseserrata in the deciduous forest.     

Gnawing damage by rodents to the seedlings ofFagus crenata andQuercus mongolica var.grosseserrata in a temperateSasa grassland-deciduous forest series in southwestern Japan

The effects of dwarf bamboo,Sasa, cover on the initial morrality of hardwood seedlings were investigated by transplanting 1-year-old beech (Fagus crenata) and current-year oak (Quercus mongolica var.grosseserrata) seedling to three different stands; old-growth beech and secondary oak forests withSasa undergrowth, and aSasa grassland in a grassland-forest series near the top of Mt Jippo, southwestern Japan. The most frequent cause of seedling morrality was gnawing of the stems by rodents. In the beech forest, the gnawing was more likely to occur underSasa cover, suggesting that it provides a good habitat for rodents on the beech forest floor. TheSasa under growth may thus play an imporrant role in regeneration of beech forest. In the oak floor, mortality of both species was low and only a little gnawing occurred during a year. However, no natural oak seedling were found in the forest even after a mast year. This may be because most of the acorns disappeated before establishment. The early-stage demography of hardwood seedling as oak may thus play an imporrant role in regeneration of oak forest. In theSasa grassland where the seed supply is small, almost all of the seedlings died fromo gnawing regardless of the presence ofSasa cover. These factors prevent the recruitment of a sizable seedling bank. Rodents may thus play an imporrant role in maintenance of theSasa grassland.     

Stand dynamics during a 12-year period in a second-growth stand in a cool temperature forest in northern Japan

In 1981 and 1993, trees over 2 m high were measured and mapped to clarify stand dynamics in two permanent plots of 0.1 ha in a secondary, cool temperate, mixed broadleaf/conifer forest after logging in the Tomakomai Experiment Forest, central Hokkaido, northern Japan. The species could be placed in two groups according to the change in basal area and density, and annual height growth:Quercus mongolica var.grosseserrata, Phellodendron amurense, Prunus sargentii andTilia japonica (Group I: GI) showed episodic regeneration and/or fast height growth;Acer mono, Acer palmatum var.matsumuae andSorbus alnifolia (Group II: GII) showed continuous regeneration and/or slow height growth. The age distribution of stems over 10 cm in d.b.h. suggests synchronous regeneration of GI and GII species. Vertical stratification was promoted during the 12-year period by the difference in annual height growth between the two species groups. Additional tree censuses in both remnant old-growth stands and second-growth stands after large scale blowdowns demonstrated thatQ. mongolica var.grosseserrata is most dominant in stands varying in the stages of development. Disturbance history and successional trends in dominant species implied that stand-devastating disturbances were responsible for the regeneration ofQ. mongolica var.grosseserrata in the study forest.     

Influence of canopy tree phenology on understorey populations of Fagus crenata

Question: Is light available for subcanopy individuals of Fagus crenata spatiotemporally heterogeneous across patches with closed canopies of different foliage phenologies and gaps? Is local abundance of Sasa influenced by the composition of the canopy layer? If so, does the Sasa layer also affect the amount of light available to small F. crenata saplings? Is variation in F. crenata population structure consistent with the hypothesis that light is important? Location: Mt. Kurikoma, Japan 780 m a.s.l. Methods: Population structure of subcanopy individuals of Fagus crenata and importance of Sasa were examined in five patch types. The patch‐types were Fc (F. crenata only in the crown), Qm (Quercus mongolica var. grosseserrata only in the crown), Mo (Magnolia obovata only in the crown), Fc’ (periphery of F. crenata) and Gap. Seasonal changes in light availability above and below the Sasa layer was examined by using hemispherical photographs and quantum sensors. Results: Subcanopy individuals of F. crenata began unfolding their leaves approximately one month earlier than canopy trees of Q. mongolica var. grosseserrata and M. obovata, but a few days later than those of adult F. crenata. Accumulated photosynthetic photon flux density above the Sasa layer was greatest in Qm and Mo, and lowest in Fc. Importance of Sasa was highest in Gap. Maximum height and the number of subcanopy individuals of F. crenata were greatest in Qm, followed by Mo, and lowest in Fc. Conclusions: Differences in canopy layer composition probably influence the regeneration of F. crenata both directly through their foliage phenologies, and also indirectly by determining the importance of Sasa.     

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.     

Shoot structure and dynamics of saplings and canopies of three deciduous broad-leaved trees of a coppice forest in central Japan

Three deciduous broad-leaved trees, Quercus serrata, Castanea crenata and Carpinus laxiflora, were the main constituents of a coppice forest in central Japan. The shoot elongation and leaf emergence modes of both saplings and the canopy of the three species were investigated. The shoot elongation modes of Q. serrata and C. crenata were the same in saplings and the upper layer of the canopy. The second shoots of these two species were formed after the first shoots were elongated. C. laxiflora was different between saplings and the upper layer of the canopy. In saplings, only the first shoots took a long time to elongate. In the upper canopy layer, higher order shoots were formed in the same way as in the other two species. In the lower layer of the canopy, all three species showed the same shoot elongation mode, in which only the first shoot and its duration of elongation was short. Leaf longevity, individual leaf area, leaf mass per unit leaf area and the stem mass per unit stem length of C. laxiflora were significantly shorter or significantly smaller than those of Q. serrata and C. crenata. The length of the stem per unit leaf area of C. laxiflora was three times that of Q. serrata and five times that of C. crenata. The elongation growth of C. laxiflora was highly efficient as it occurred with a small leaf area. The shoot dynamics and the shoot structure of C. laxiflora are more suitable for elongation growth than in Q. serrata and C. crenata. Furthermore, the shoot structures of the three species were compared and ecological characteristics of the three species are discussed. Received: 29 September 1998 / Accepted: 17 September 1999     

Early post-fire vegetation regeneration in <Emphasis Type="Italic">Larix kaempferi</Emphasis> artificial forests with an undergrowth of <Emphasis Type="Italic">Sasa senanensis</Emphasis>

Post-fire vegetation regeneration was studied for a 6-year period in a 13-year-old-artificial forest consisting of Larix kaempferi with a dense undergrowth of Sasa senanensis. The study site was classified into three fire severity categories according to the degree of Sasa senanensis scorching, that is, a high-severity category, a mid-severity category, and a low-severity category. Study plots were established in areas which fitted the criteria for each category, and in nearby unburned sites. A total of 41 woody species were newly emerged during the 6-year study period in the burned and unburned plots. Only a few seedlings and resprouts emerged in the unburned plots, while many seedlings emerged in the high-severity plots from the first year after fire onward. A high-severity fire that burns the rhizomes of Sasa is necessary for the vegetation recovery by germination of seed. Whereas the establishment of seedlings was restricted to a few years after fire, the regeneration through resprouting continued into the last year of observation. The survival time of resprouts was longer than that of seedlings, and the survival time of shade-tolerant species was longer than that of shade-intolerant species. In contrast, shade-intolerant species grew more rapidly than shade-tolerant species. The plants ability to exceed the maximum height of the Sasa before the bamboo recovers can be critical to the survival of shade-intolerant species. Because resprouts have a stronger resistance to the shade of Sasa than seedlings, the resprouts of shade-tolerant species play a major role in the re-establishment of woody species after fire in sites with considerable Sasa ground-cover.     

Nutrient compositions of acorns and horse chestnuts in relation to seed-hoarding

Nutritional compositions of acorns (Quercus serrata Thunb. and Quercus mongolica Fisch. var. grosseserrata Rehd. et Wils.) and horse chestnuts (Aesculus turbinata Blume) were analyzed. Major nutrients of acorns and horse chestnuts were carbohydrate, protein and fat. They proved to contain considerable amounts of tannins (7.28–11.72% dry mass^–1) and saponins (6.20%), respectively. The nutrients and the secondary metabolites of seeds buried for either 1 month or 3 months were analyzed to evaluate effects of seed-caching by animals. Noticeable changes were not observed, which suggested that caching may not bring any advantages to the seed-hoarders with respect to nutrition.     

Regeneration of beech (Fagus crenata) after the simultaneous death of undergrowing dwarf bamboo (Sasa kurilensis)

Seedling recruitment and survivorship of beech (Fagus crenata) were studied with special reference to the simultaneous death of undergrowing bamboo (Sasa kurilensis). The survival rate of beech seedlings on the floor whereSasa had withered was much higher than that on the floor whereSasa survived. Damping off caused the largest mortality among beech seedlings. However, the allocation pattern of matter to different parts of the seedlings indicated that their survival was greatly affected by production economy. The dense cover of dwarf bamboo prevented the establishment of beech seedling banks on the forest floor. The interval between the times when simultaneous death ofSasa occur and the length of its recovery period are thus important factors controlling the dynamics of beech forests in Japan.     

Unveiling the origin of Quercus serrata subsp. mongolicoides found in Honshu,Japan, by using genetic and morphological analyses

Quercus mongolica is a tree found in temperate deciduous forests in east Asia. In Japan, Q. mongolica var. crispula is commonly found; moreover, an oak whose morphology is similar to that of Q. mongolica var. mongolica of the Asian continent has been found in certain areas of Honshu and Hokkaido. Recently, the oak found in Honshu was described as Q. serrata subsp. mongolicoides (QSM). However, genetic comparison between this oak and Q. mongolica var. mongolica of the Asian continent has not been performed; the origin of QSM is thus unknown. Therefore, we aimed to determine the origin of QSM by conducting nuclear microsatellite (nSSR), chloroplast DNA (cpDNA) and leaf morphology analyses for the three taxa, as well as other congeners. The cpDNA variation overlapped among the three taxa, suggesting low discrimination ability for these taxa. Although morphological congruency was found between QSM and Q. mongolica var. mongolica, results of nSSR analyses showed that QSM contained a genetic admixture of Q. mongolica var. mongolica of the Asian continent and Q. mongolica var. crispula of Japan, bolstering an admixture hypothesis. The nSSR and cpDNA analyses suggested that Q. mongolica var. crispula can be the progenitor of Q. mongolica var. mongolica and harbors the ancestral cpDNA haplotypes. Therefore, we concluded that QSM might have been created by an admixture that likely occurred within Japan between Q. mongolica var. crispula and putative relict Q. mongolica var. mongolica, which might have diverged in or around Japan from Q. mongolica var. crispula during the late Pleistocene.     

Fallen leaves and unpalatable plants as alternative foods for sika deer under food limitation

Feeding behavior and rumen contents of sika deer (Cervus nippon Temminck) under food limitation were studied on Nakanoshima Island, Hokkaido. During the phase of population growth, the deer subsisted on tree bark and twigs, deciduous leaves and dwarf bamboos (Sasa spp.) in the winter. After a crash in population, the deer began feeding on the fallen leaves of deciduous trees and continued to do so throughout the year. They also ate unpalatable plants Cephalotaxus harringtonia var. nana Rehd., Senecio cannabifolius Less. and Cynanchum caudatum Maxim. as winter foods, which used to remain untouched by deer, and had accordingly expanded their distributions, following a decrease in the amount of dwarf bamboos available. These facts suggest that sika deer drastically shift their foods and exploit alternative foods under conditions of food limitation.     

Importance of seed size for the establishment of seedlings of five deciduous broad-leaved tree species

Effects of seed size and phenology on the establishment of five deciduous broad-leaved tree species were examined in deciduous woodland. Treatments included absence and presence of litter in the forest understory, a small gap, and a large gap. Seedling emergence of large-seeded speciesQuercus mongolica var.grosseserrata andAcer mono was not reduced by accumulation of litter in the forest understory, but was promoted in the large gap where litter was less. Seedling emergence of small-seeded species,Alnus hirsuta, Cercidiphyllum japonicum andBetula platyphylla var.japonica, was reduced by the litter in almost all of the sites. Seedlings of large-seeded species avoid shade stress phenologically by unfolding all of their large leaves in a short period before canopy closure in the forest understory. These species had little mortality after seedling emergence. In contrast, small-seeded species have a longer duration of leaf emergence, shorter leaf longevity, and rapid leaf turnover in all the sites. These seedlings attained similar height to those of the large-seeded species at the end of the second year in the large gap, but survival and height growth rate decreased after canopy closure in the forest understory. We suggest that the importance of seed size in determining seedling establishment largely depends on the relationships between seasonal changes of environmental conditions and phenological traits of seedlings, which are related to seed size.Abbreviations Ah Alnus hirsuta - Am Acer mono - Cj Cercidiphyllum japonicum - Bp Betula platyphylla var.japonica - Qm Quercus mongolica var.grosseserrata     

A phytosociological analysis of woody species in forest communities of a part of Kumaun Himalaya

This paper reports on a detailed phytosociological analysis of forests in the NW catchment of the Gola River in Kumaun Himalaya, 29°19–29°27N and 79°32–79°42E. Fourteen sites and 56 stands at elevations ranging from 1200 to 2523 m and covering the following five forest types were investigated: Pinus roxburghii, mixed, Quercus leucotrichophora, Q. lanuginosa, and Q. floribunda. The basal cover of the forests differed according to slope position and aspect. The three oak forests had more basal cover than the other two, and Q. lanuginosa had the most. The performance of individual tree and shrub species and the number of saplings and seedlings differed according to slope position and aspect. The mixed forest had the greatest tree diversity, and among the others diversity increased with increasing basal cover. The diversity of trees, saplings, and herb layer was greatest on aspects with intermediate temperature and moisture conditions; whereas that of shrubs and seedlings increased towards the cooler (and wetter) and warmer (and drier) exposures. There was a positive relation between the diversity of shrubs plus seedlings and trees plus saplings in P. roxburghii and mixed forests; whereas this relationship was inverse in the three oak forests. In general, the dominance-diversity curves for the tree layer followed a geometric series conforming to the niche pre-emption situation in communities of low diversity. Among the forests, the regeneration was best in Q. lanuginosa and worst in Q. leucotrichophora.Nomenclature follows: Osmaston (1926).Finacial support from the Council of Scientific and Industrial Research, New Delhi and the Indian Space Research Organisation, Banglore is gratefully acknowledged.     

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.     

Herbivory pattern and induced responses to simulated herbivory inQuercus mongolica var.grosseserrata

Seasonal changes in leaf traits and the herbivory pattern ofQuercus mongolica var.grosseserrata were studied, and simulated herbivory experiments were carried out in order to evaluate leaf trait responses. Leaves ofQ. mongolica emerged simultaneously in spring and most were retained until autumn. Nitrogen concentration was highest when leaves first emerged and decreased rapidly with leaf age. Leaf mass per area (LMA) increased with leaf age. Herbivore attack was concentrated in the first 20 days after bud-break, which corresponded to the high nutritional value of the leaves for herbivores at this time. Simulated herbivory experiments indicated that LMA increased with artificial leaf damage, suggesting an increase in leaf toughness, and that nitrogen concentration decreased later in the season in comparison with intact leaves. As a result, herbivore attack following artificial leaf damage decreased with increasing initial leaf damage. However, leaf longevity was not affected by initial leaf damage. These responses were considered to be a strategy to disperse herbivory damage among leaves.     

Effects of understory dwarf bamboo on soil water and the growth of overstory trees in a dense secondary <Emphasis Type="Italic">Betula ermanii</Emphasis> forest,northern Japan

The effects of understory dwarf bamboo (Sasa kurilensis) on soil water and the growth of overstory trees were studied in a dense secondary forest of Betula ermanii in northern Japan. Four plots were established in a Betula ermanii forest with Sasa kurilensis in the understory. The Sasa was removed in two of the plots. The annual increment of the trunk diameter for each tree was measured in the first two years from the commencement of the experiment. Soil water potential was similar in the plots following significant rainfall, but was found to be greater in the plot without Sasa between rainfall events. This suggests that the removal of Sasa slows the reduction of soil water after rainfall. The relative growth rate of the trunk diameter of Betula ermanii increased with tree size in all of the plots because taller trees strongly suppressed smaller ones in the dense forest. The growth rates of Betula ermanii were higher in the plots without Sasa. However, the difference in growth rates between all of the plots tended to be smaller in smaller size classes, possibly because smaller trees were strongly suppressed by larger ones, irrespective of the presence/absence of Sasa. Therefore, the removal of Sasa increased soil water and encouraged the growth of larger Betula ermanii in dense forest during the first two years after the Sasa was removed. The present study suggests that Sasa can reduce the growth of larger Betula ermanii in dense forest by limiting available soil water to these trees.     

A newOphiostoma species isolated from Japanese oak infested byPlatypus quercivorus

A newOphiostoma species was isolated from the sapwood ofQuercus mongolica var.grosseserrata and described asOphiostoma longicollum sp. nov. The species is characterized by a long perithecial neck with a gelatinous cap at the apex and a constricted base, orange-section-shaped ascospores that accumulate in a yellow-orange drop at the apex of the neck, and itsSporothrix anamorph. Contribution No. 137, Laboratory of Plant Pathology and Mycology, Institute of Agriculture and Forestry, University of Tsukuba.     

Early seedling growth of pine (Pinus densiflora) and oaks (Quercus serrata,Q. mongolica,Q. variabilis) in response to light intensity and soil moisture

In climatic chambers seed germination and seedling growth of Pinus densiflora Sieb. et Zucc., Quercus serrata Thunb., Quercus mongolica Fisch. ex Turcz. and Quercus variablilis Bl. were investigated as functions of light intensity and soil moisture. In Korea these tree species occur widely and form mixed forests with different distributions. Species clearly differed in the pattern of germination and early seedling growth between light and soil treatments. The germination of pine did not differ between the experimental treatments until the breaking of the primary buds. After that, light intensity was the deciding factor for further development. In the most moist treatment (approx. field capacity) growth of the pine seedlings was strongly inhibited. For the three oak species, differences between experimental treatments first occurred after complete formation of primary leaves. Seed development strongly correlated with the weight of the acorn. The large seeded Q. variabilis (acorns with mean weight of 4.7 g) developed faster and reached larger dimensions towards the end of the experiment than Q. mongolica (2.8 g per acorn) and Q. serrata (0.9 g per acorn). Regarding height and biomass growth, the oak species showed a higher shade tolerance than pine. The proleptic shoot growth was clearly influenced by the light intensity. Root formation was favoured by a high exposure to light. In case of the oak species reduction of soil moisture increased the length of primary roots and the number of secondary roots.     

Dynamics and conservation of the gwangneung national forest in central Korea: A national model for forest restoration

Forests in Gwangneung National Arboretum District (GNAD) have been protected since the 15^th century. Consequently, these forests support more than 20% of all plant species in Korea. We constructed vegetation maps for landscape analysis, and forest dynamics, species diversity, and sustainable management were discussed. Secondary forests compose 51.0% of the whole vegetation, while plantations compose 45.2%.Quercus serrata dominates the forest, and the plantations are comprised mainly ofPinus koraiensis. Although dominated by plantations and human installations, the presence of a rare riparian hardwood forest, composed primarily of Q.aliena, was notable. Species diversity of the riparian (H′ = 3.38) was significantly (p<0.0001) higher than the upland (H′ = 1.56). Species turnover rate as a spatial heterogeneity was also higher. Such high species diversity and heterogeneity are justification to conserve the riparian and lowland forests in GNAD. Extensive recruitments of their own seedlings and saplings suggest a sustainable regeneration of Q.serrata and Q.aliena stands in the lower elevations, and the opposite is true for the Q.mongolica andP. rigida stands in higher elevations. GNAD contains diverse natural landscape elements that range from riparian to upland vegetation, which may well serve as a national model for forest restoration.