Pre- and post-dispersal damage to the nuts of two beech species (Fagus crenata blume andF. japonica Maxim.) that masted simultaneously at the same site

A study of pre- and post-dispersal damage to the nuts of two beeches,Fagus crenata Blume andF. japonica Maxim., which masted simultaneously, was performed on a single site in Japan. Most insect damage to the nuts before dispersal resulted from boring by lepidopteran larvae, and the level of attack was markedly higher onF. crenata than onF. japonica. Early nut growth ofF. crenata allowed these borers to act as the first attackers on the nuts. Conversely, late nut growth ofF. japonica delayed the attack of borers, and foliage feeders acted as the first attackers instead, chewing on nuts from the outside. The low level of borer attacks on the nuts ofF. japonica in mast years was thought to have been caused by foliage feeders decreasing the amount of food resource available for later attack by borers in lean years. This could then lead to a subsequent depression of the population of borers in mast years. The proportion of nuts damaged by moth larvae was independent of the density of nuts for both beech species. Caching by vertebrates was thought to be the most important factor in post-dispersal damage because the density of nuts of both beech species on the forest floor in March was half the level measured using seed traps in the previous year. However, post-germination damage caused by a fungus was also an important factor. The survival rate of the nuts (including current-year seedlings) in the year after dispersal was independent of the density of dispersed sound nuts in the previous year for both beech species, with the exception of one set of observations (forF. japonica in July). This research was supported in part by a grant from the Japanese Ministry of Agriculture, Forestry and Fisheries.     

Notes on beech (Fagus crenata Blume) seed and seedling mortality due to rodent herbivory in a northernmost beech forest, Utasai, Hokkaido

We investigated the mortality of post-dispersal seeds and current-year seedlings of a beech species (Fagus crenata Blume) at the northernmost extent of its geographical range at Utasai (Kuromatsunai), Hokkaido, Japan. Introduced seeds that were protected from rodent predation by wire cages had a higher survival rate from winter to the following spring (mean ± SD: 84 ± 16.7%,n = 10) than controls without cages (mean ± SD: 12 ± 17.9%,n = 10). Unprotected transplanted current-year seedlings with almost fully opened primary leaves never survived more than three days (0%,n = 30), while more than 80% (n = 30) of seedlings within cages survived to the end of the growing season. These results indicate that rodent herbivory is a major mortality factor that strongly prevents the establishment of beech seedlings. In a natural population, we observed that 90% of seedlings (n = 197) disappeared within 10 days after their emergence and rodent herbivory caused this heavy mortality. A herbivorous rodent, the gray-backed vole,Clethrionomys rufocanus bedfordiae, was remarkably abundant in late June (101.5 voles/ha), suggesting that this species strongly affects the formation of the seedling bank. The presence ofC. rufocanus bedfordiae may be one of the reasons for the scarcity of beech seedlings and saplings and the rarity of recruitment in this northernmost beech forest.     

Induced response of the Siebold's beech (Fagus crenata Blume) to manual defoliation

Siebold's beech (Fagus crenata) was manually defoliated for two successive years. The beech caterpillar (Quadricalcarifera punctatella) was used in a bioassay to determine insect performance. Survival and body size were low on foliage from defoliated trees. Reduced foliar nitrogen and increased tannin content were probably the main causes of the low insect performance. Leaves were less tough on defoliated trees than in controls. Two sucessive years of manual defoliation caused stronger induced resistance than one year defoliation. The quality, as well as the quality of the foliage, decreased the year following manual defoliation; total weight of leaves on a tree was less than one half of that before treatment. Severe defoliation may cause a decrease of leaves the following year and starvation may limit populations. Delayed induced resistance of beech trees is proposed as a possible cause of the cyclical population dynamics ofQ. punctatella. The delayed induced response also affected folivorous insects other thanQ. punctatella.     

Spatial and seasonal variations in photosynthetic properties within a beech (Fagus crenata Blume) crown

The light response curve and the intercellular CO₂ concentration response curve of CO₂ assimilation rate were investigated together with the light conditions at the four different heights within the beech crown from 1995 to 1997 on Mt. Fuji in Japan. On the seasonal fluctuation, the CO₂ assimilation rate at light saturated condition increased rapidly in May, and attained to the maximum between the end of June and July, thereafter, slightly decreased until the middle of August and rapidly decreased in September and October. The daily sum of photosynthetic photon flux density attenuated with deeping within the crown, and particularly, the relative value on 2nd position dropped to only 30%. TheA _max decreased from 10 to 5μmol m⁻² s⁻¹, approximately, with deeping within the crown. The light saturation point, quantum yield, light compensation point and dark respiration rate also varied with deeping. These results suggest that the photosynthetic properties vary gradually from sun to shade leaves along the light attenuation within a beech crown. At light saturated condition, the stomatal conductance and mesophyll conductance were strongly correlated withA _max among the four different heights (r > 0.96, respectively). TheC _i/C _a ratio was around 0.8, and there were no remarkable differences among the four different heights. These results suggest that the vertical gradient ofA _max depends on the variation of mesophyll conductance. The stomatal conductance may be also one of the major factor in the vertical gradient ofA _max. However the intercellular CO₂ concentration doesn’t influence the vertical gradient ofA _max within the crown. This work is supported by the Sasagawa Scientific Research Grant from The Japan Science Society and Grant-in-Aid for Scientific Research (C).     

Effects of root trenching of overstorey Norway spruce (Piceaabies) on growth and biomass of underplanted beech (Fagussylvatica) and Douglas fir (Pseudotsugamenziesii) saplings

In Central Europe, the conversion of pure Norway spruce stands (Picea abies [L.] Karst.) into mixed stands with beech (Fagus silvatica L.) and other species like e.g. Douglas fir (Pseudotsuga menziesii [Mirb.] Franco) is accomplished mainly by underplanting of seedlings beneath the canopy of overstorey spruce trees after partial cutting treatments what means exposure to shade and below-ground root competition by the overstorey to the seedlings. Particularly about the second factor, our knowledge is limited. Therefore, we carried out a below-ground competition exclusion experiment by root trenching and investigated the effects on soil resources, growth, and biomass partitioning of underplanted beech and Douglas fir saplings under target diameter and strip cutting treatments. The exclusion of overstorey root competition by trenching increased the soil water potential in the second year that had a fairly dry growing season and led to significantly higher foliar concentrations of most nutrients, particularly in Douglas fir, indicating an amended nutrient supply. Both improvements were accompanied by an increase in length and diameter increment of the underplanted saplings, appearing in both species only after having surpassed a species-specific threshold light value (Douglas fir 16% of above canopy radiation, beech 22%). We also found significant interactions between trenching and light for specific fine root length and further biomass and morphological parameters. Judged by the much steeper increase in height and diameter growth with increasing light after release from below-ground competition, Douglas fir saplings appeared to be more sensitive to root competition than beech saplings what conforms to older findings for beech. According to our results, a strip cutting seems to be more appropriate than a target diameter cutting treatment to replace a pure spruce stand by a mixed stand with beech and Douglas fir.     

Canopy gaps and regeneration in old-growth Oriental beech (Fagus orientalis Lipsky) stands, northern Iran

Virgin beech Fagus orientalis forests in northern Iran provide a unique opportunity to study the disturbance regimes of forest ecosystems without human influence. The aim of this research was to describe characteristics of natural canopy gaps and gap area fraction as an environmental influence on the success of beech seedling establishment in mature beech stands. All canopy gaps and related forest parameters were measured within three 25 ha areas within the Gorazbon compartment of the University of Tehran’s Kheyrud Experimental Forest. An average of 3 gaps/ha occurred in the forest and gap sizes ranged from 19 to 1250 m² in size. The most frequent (58%) canopy gaps were <200 m². In total, canopy gaps covered 9.3% of the forest area. Gaps <400 m² in size were irregular in shape, but larger gaps did not differ significantly in shape from a circle. Most gaps (41%) were formed by a single tree-fall event and beech made up 63% of gap makers and 93% of gap fillers. Frequency and diversity of tree seedlings were not significantly correlated with gap size. The minimum gap size that contained at least one beech gap-filling sapling (<1.3 m tall) was 23.7 m². The median gap size containing at least one beech gap-filling sapling was 206 m² and the maximum size was 1808 m². The management implications from our study suggest that the creation of small and medium sized gaps in mixed beech forest should mimic natural disturbance regimes and provide suitable conditions for successful beech regeneration.     

Response of oriental beech (Fagus orientalis Lipsky) seedlings to canopy gap size

The response of nursery seedlings and wildings of oriental beech(Fagus orientalis Lipsky) to canopy gap size was studied ina climax beech forest in northern Iran with respect to seedlingsurvival, shoot growth and vitality. These parameters were followedfor 2 years after planting in gaps of 50, 200 and 600 m², aswell as in the open field. Foliage coloration was used as acriterion for vitality. The survival rate of nursery seedlingsand wildings dropped rapidly with increasing gap size from 84per cent in 50-m² canopy openings to     

Japanese beech (Fagus crenata) plantations established from seedlings of non-native genetic lineages

In order to examine whether seedlings of non-native genetic lineages had been planted, we compared chloroplast DNA haplotypes in plantations and neighboring natural populations of Fagus crenata in Nagano Prefecture, Japan. This region was chosen for study because there are abundant natural F. crenata populations with different haplotypes, and because the species is also frequently used for afforestation in the area. We sampled 159 trees from 30 populations of the species across most of its natural range in the region, and 136 trees from 20 plantations of the species, ranging in age from 3 to 17 years. Six single nucleotide polymorphisms and four insertions/deletions in two chloroplast DNA regions (trnL–trnF and trnK) were analyzed in each of the sampled individuals in order to determine their haplotypes. Four haplotypes (B, D, E and F) were detected in the natural populations, and these exhibited a clear geographical structure; in contrast, only two haplotypes (A and B) were found in the plantations. We found that 15 out of the 20 plantations—located in central and southern areas on the Pacific side of Honshu, where the natural populations contain haplotypes D, E, and F, and where the climate is characterized by dry, cold winters—had been established using seedlings with haplotypes A and B derived from the Sea of Japan side of the Japanese Islands, where it snows heavily. The risks associated with planting seedlings of inappropriate lineages are discussed.     

Canopy recruitment in the beech (<Emphasis Type="Italic">Fagus engleriana</Emphasis>) forest of Mt. Shennongjia,Central China

A dendrochronological technique was used to investigate canopy recruitment of beech (Fagus engleriana Seem.) trees in western Hubei Province of Central China by identifying growth releases through gap formation. The results indicated that 83% and 17% of the canopy trees regenerate in gaps and the understory, respectively. Radial growth analyses demonstrated that there are three modes for beech regeneration and canopy recruitment: (1) gap origin without suppression; (2) gap origin with period(s) of suppression; and (3) understory origin. Multiple periods of release and suppression imply that beech regeneration and canopy recruitment are very sensitive to gap formation.     

Beech leaf disease symptoms caused by newly recognized nematode subspecies Litylenchus crenatae mccannii (Anguinata) described from Fagus grandifolia in North America

Symptoms of beech leaf disease (BLD), first reported in Ohio in 2012, include interveinal greening, thickening and often chlorosis in leaves, canopy thinning and mortality. Nematodes from diseased leaves of American beech (Fagus grandifolia) sent by the Ohio Department of Agriculture to the USDA, Beltsville, MD in autumn 2017 were identified as the first recorded North American population of Litylenchus crenatae (Nematology, 21, 2019, 5), originally described from Japan. This and other populations from Ohio, Pennsylvania and the neighbouring province of Ontario, Canada showed some differences in morphometric averages among females compared to the Japanese population. Ribosomal DNA marker sequences were nearly identical to the population from Japan. A sequence for the COI marker was also generated, although it was not available from the Japanese population. The nematode was not encountered in Fagus crenata (its host in Japan) living among nematode‐infested Fagus grandifolia in the Holden Arboretum, nor has L. crenatae been reported in American beech in Japan. The morphological and host range differences in North American populations are nomenclaturally distinguished as L. crenatae mccannii ssp. n. from the population in Japan. Low‐temperature scanning electron microscopy (LT‐SEM) demonstrated five lip annules and a highly flexible cuticle. Females, juveniles and eggs were imaged within buds with a Hirox Digital microscope and an LT‐SEM. Nematodes swarmed to the tips of freshly cut beech buds, but explants could not be maintained. Inoculation of fresh nematodes from infested leaves or buds to buds or leaves of F. grandifolia seedlings resulted in BLD leaf symptoms. Injuring dormant buds prior to nematode application, in fall or spring, promoted the most reliable symptom expression. The biogeography and physiology of anguinid nematode leaf galling, and potential co‐factors and transmission are discussed.     

Pathogenicity of Colletotrichum dematium isolated from current-year beech seedlings exhibiting damping-off

The pathogenicity of Colletotrichum dematium was evaluated as the agent causing post-emergence damping-off of current-year beech (Fagus crenata) seedlings. Current-year beech seedlings inoculated with C. dematium died within 20 days after both wound and non-wound inoculation, indicating that this fungus is pathogenic to current-year seedlings. C. dematium was frequently isolated from hypocotyls exhibiting damping-off at five different sites. The incidence of isolation was high and little variation occurred between sites and years. C. dematium was also detected in all litter samples examined although the incidence of detection differed depending on sites and/or detection methods. These results strongly indicate that C. dematium plays an important role in rapid seedling death during the early stages of beech regeneration.     

Temporal and spatial dynamics of an old-growth beech forest in western Japan

Dendrochronological approaches enable us to understand forest stand dynamics by estimation of disturbance history and age structure. The present study was conducted in an old-growth beech forest in a forest reserve in western Japan. Increment cores were taken for tree ring analysis from all canopy trees in a 50 m × 130 m study plot. Radial growth release criteria were developed to identify significant growth releases in each tree ring series and to characterize the disturbance history of the study site. The age structure of the forest was indicative of continuous establishment by Fagus crenata and simultaneous establishment by Magnolia obovata. A variety of low-intensity disturbances were identified in each decade, especially after the 1900s, but the occurrence of high-intensity catastrophic disturbance was rare, and likely played an important role in maintaining species diversity in the existing forest canopy. The results also suggest that F. crenata regenerates gradually before and after both large- and small-scale disturbances, whereas M. obovata and Betula grossa regenerate only after large-scale catastrophic disturbances.     

Age structure of wind-felled canopy trees for Siebold’s beech (<Emphasis Type="Italic">Fagus crenata</Emphasis>) in the northernmost population in Karibayama,Hokkaido

To determine the lifespan of Siebold’s beech (Fagus crenata), a dominant canopy tree species of the cool temperate forests of Japan, we estimated the age of 33 mature beech trees felled by a large typhoon in September 2004 at Karibayama, in southern Hokkaido. The study site lies within one of the northernmost distributions of Siebold’s beech populations in Japan. We counted tree rings from tree disks at a height of 1 m, and diameter growth was measured in annual increments. The sample age revealed two major peaks at 240 and 175 years. The diameter growth was good between 40 and 60 years of age and gradually decreased after 70 years of age. Annual diameter increments were noticeably small in good fruiting years, indicating a large allocation of annual carbon gains to the reproductive organs. Bud scale counts of juvenile trees revealed that the age at which the height reached 1 m was approximately 6–13 years. The lifespan of Siebold’s beech in the Karibayama population was therefore estimated to be approximately 250 years, an estimation that is higher than that previously reported for this species in the northernmost populations.     

Effect of canopy gap size and ecological factors on species diversity and beech seedlings in managed beech stands in Hyrcanian forests

We studied the species diversity of the herb layer and ecological factors in harvest-created gaps in beech stands under a single-tree selection system in Northern Iran. To determine diversity, the number of beech seedlings, and other ecological factors, 16 gaps were selected and subplots of 5 m2 were positioned at the centre and at the cardinal points of each gap. Species richness and Simpson diversity index increased with increasing gap area as did numbers of seedlings. With increasing humus layer thickness, species richness declined but the Hill evenness index increased. Species richness increased with increasing light availability. There was no relationship between crown radii of beech trees and diversity indices. Correlations between environmental factors and numbers ofindividuals of some species in the herb layer were not significant except in a few cases. The results help explain the effects of man-made gaps on the dynamics of managed beech stands and this benefits evaluation of silvicultural operating plans.     

Polymorphism analysis of Fagaceae and DNA-based identification ofFagus species grown in Japan based on therbcL gene

Fagaceae species in Japan were identified by restriction fragment length polymorphism (RFLP) and sequence comparison of a region ofrbcL. Of nine restriction endonucleases used for digestion, three (MspI,RsaI,HaeIII) produced different restriction patterns in Fagaceae. Digestion byMspI yielded four patterns: Fagus species,Castanea crenata, Pasania glabra, and others. Digestion byRsaI andHaeIII afforded two patterns:Fagus species and others. These facts indicate thatCastanea crenata andPasania glabra can be identified byMspI restriction patterns ofrbcL. Sequence comparison of a region of therbcL gene among 20 species of Fagaceae showed that: (1) they could be divided into seven groups; (2) there is a site mutation betweenFagus crenata andF. japonica. The latter indicates that the wood of both Fagus species are identifiable at the species level, which is not the case using conventional methods. This result indicates the possibility of wood identification based on DNA polymorphism in Fagaceae at the intrageneric level.Part of this paper was presented at the 46th annual meeting of the Japan Wood Research Society, Kumamoto, April 3–5, 1996 and the 47th annual meeting of the Japan Wood Research Society, Kochi, April 3–5, 1997     

Photosynthetic responses to lightflecks ofFagus crenata seedlings grown in a gap and understory of a deciduous forest

Photosynthetic responses to a series of 1-min lightflecks (1,000μmol m⁻² s⁻¹) superimposed on a background with different duration (1, 5, and 10 min) and intensity (25 and 50μmol m⁻² s⁻¹) of low background photosynthetic photon flux density (PPFD) were measured in the leaves ofFagus crenata grown in a gap and understory of aFagus crenata forest in the Naeba Mountains. The two background PPFD intensities most frequently occurred in understory and gap sites respectively. The maximum net photosynthetic rate (P _Nmax) and maximum stomatal conductance (g _smax) were higher in the gap seedlings than in the understory seedlings. However, when the background PPFD was 25μmol m⁻²s⁻¹, the net photosynthetic rate (P ₂₅) and stomatal conductance (g _s25) were almost the same between the gap and understory. When the background PPFD duration was 1-min, the net photosynthetic rate (P _N ) at the end of each lightfleck increased progressively. When the background PPFD duration was 5- and 10-min, the increase inP _N at the end of each lightfleck was less. This indicates that background PPFD duration is important to photosynthetic responses to lightflecks. The higher ratios ofP ₂₅/P _Nmax andg _s25/g _smax in the understory seedlings indicate that the understory seedlings can maintain relatively lower levels of biochemical and stomatal limitations than the gap seedlings under low light conditions. The ratios ofP _N /P _Nmax at the end of each lightfleck (IS) and light utilization efficiency of single lightflecks (LUE _s) that showed the influence of lightflecks on carbon gain were higher in the understory seedlings than in the gap seedlings when the background PPFD was 25μmol m⁻² s⁻¹. This means that understory seedling are capable of utilizing fluctuating light more efficiently under low light conditions than the gap seedlings although the net carbon gain of single lightflecks (CG _s) in the understory seedlings was not higher than that in the gap seedlings. There were no significant differences inIS andLUE _s between understory seedlings at a background PPFD of 25μmol m⁻² s⁻¹ and gap seedlings at a background PPFD of 50μmol m⁻² s⁻¹. However,CG _s in gap seedlings was higher than in understory seedlings. These results provide more evidence thatF. crenata acclimate to a natural light environment in respect to relative induction state at low background PPFD and can capture the fluctuating light at the same efficiency in both the gap and understory seedlings under natural light environments. This study was funded by the research project, Evaluation of Total CO₂ Budget in Forest Ecosystems, coordinated by the Ministry of Agriculture, Forestry and Fisheries of Japan.     

Gap dynamics in the Western Carpathian mixed beech old-growth forests affected by spruce bark beetle outbreak

Understanding how species-specific disturbances affect the dynamics of mixed forests is becoming increasingly important due to rapidly changing disturbance regimes. This study estimated the effect of Norway spruce (Picea abies (L.) Karst.) mortality on the disturbance processes in two mixed beech stands of the Western Carpathians that were affected by a bark beetle outbreak. We evaluated the size distribution, fraction of canopy and expanded gaps, the characteristics of gapmakers and the contribution of different species to gap size. The measured canopy gap fraction was <5%, and most canopy gaps were small (<100 m²). Spruce was the most abundant gapmaker, and its share among gapmakers was 3–6 times higher than its share in the canopy. We found that the increase in spruce mortality due to the outbreak resulted in a fine-scale mortality pattern. However, spruce gapmakers did not contribute much to gap area size, as shown by a weak correlation between the number of spruce gapmakers and the area of expanded gaps. Diameter distribution of living versus recently dead trees showed that beech mortality occurred disproportionately in large size classes. However, dead spruce trees were equally frequent in all diameter classes, which means beetles did not exclusively attack larger trees in these stands during the outbreak. We conclude that spruce mortality may have influenced successional processes by giving a competitive advantage to two other species that were not affected by the outbreak, provided that a high deer browsing intensity does not hinder the regeneration of new seedlings.

     

Growth Dynamics During Canopy Recruitment of Sprout-Origin Stems in Japanese Beech (Fagus japonicaMaxim.) Stools in Old Growth Forests of Central Japan

Abstract

In order to clarify mechanisms of canopy recruitment of Fagus japonicastems of sprout origin, age structure and growth dynamics were studied for stems within beech stools in two stands of old growth forests on the Pacific side of central Japan. The DBH vs. height and age relationships of the beech stems showed continuous distribution from small-young, understory stems to large-old, canopy ones. There were one to seven definable periods of suppression prior to canopy recruitment in F japonica.Prior to canopy recruitment number and length of release episodes were fewer and shorter than those of suppression. E japonicastems of sprout origin can endure suppression for a long time. Continuous emergence and the alternation of fast and slow growth in F. japonicastems suggest the sprouting ability for the beech canopy can play an important role in the successful closure of canopy openings. Canopy stems showed patterns of suppression and release that were often synchronous with other canopy stems within the same beech stool. Minor disturbances such as single crown falls can play an important role in the maintenance of the multi-stem structure of beech stools in old growth forests.     

Influence of foliar phenology and shoot inclination on annual photosynthetic gain in individual beech saplings: A functional–structural modeling approach

We developed a functional–structural plant model for Fagus crenata saplings and calculated annual photosynthetic gains to determine the influences of foliar phenology and shoot inclination on the carbon economy of saplings. The model regenerated the three-dimensional shoot structure and spatial and temporal display of leaves; we calculated the hourly light interception of each leaf with a detailed light model that allowed us to estimate hourly leaf photosynthetic gain taking leaf age into account. To evaluate the importance of simultaneous foliar phenology and slanting shoots in beech saplings, we calculated the photosynthetic budgets for saplings with contrasting foliar phenologies and shoot inclinations. In our simulations, we distinguished between simultaneous and successive foliar phenologies, upright and slanting shoot inclinations, and environments with and without a vertical gradient in light intensity. Other model parameters (including photosynthesis vs. light curve, leaf size, and leaf shape) were obtained directly from live beech saplings. With no vertical gradient in light intensity, modeled saplings with simultaneous foliar phenology and slanting shoots (as in live beech) had larger annual photosynthetic gains than saplings with other combinations of traits. Hence, simultaneous foliar phenology and slanting shoots are efficient ways to display leaves in the shaded forest understory light regime where beech saplings thrive. In the presence of vertical light gradients, which can occur in canopy gaps, saplings with upright shoots had larger annual photosynthetic gains than counterparts with slanting shoots. Although mean daily photosynthetic gains of saplings with successive foliar phenology were elevated by exposing leaves to strong light when young and productive, the annual photosynthetic budget of these saplings was reduced (compared to saplings with simultaneous foliar phenology) by their relatively short leaf lifespan. Overall, our results suggest that slanting shoots with simultaneous foliar phenology are particularly successful in shaded environments, where beech often dominates, because they appear to maximize the annual carbon budget by avoiding self-shading and extending leaf lifespans.     

The effects of canopy gaps on seedling emergence and growth of two deciduous broad-leaved species (Cornus controversa andPrunus grayana) in aCryptomeria japonica plantations

The effects of canopy gaps on seedling emergence and growth ofCornus controversa andPrunus grayana were studied in a 21-year-oldCryptomeria japonica plantation. The seeds of the two species were sown in December 1995 and their fate was followed until March 2000.P. grayana germinated in 1996, butC. controversa germinated in 1997. In both species studied, more than 70% of the seedlings survived in the forest edge until the end of the experiment, although none survived in the forest understory in the first growing season. In the gap, the survival rate was higher inP. grayana than inC. controversa. In this experiment, some trees were cut to enlarge the area of the gap, in which the growth rate increased markedly forC. controversa seedlings, but not forP. grayana seedlings after the cutting. These two species showed substantial differences in the patterns of seedling emergence, survival and growth in aCr. japonica plantation which had a canopy gap.