Effects of local biotic neighbors and habitat heterogeneity on tree and shrub seedling survival in an old-growth temperate forest

Seedling dynamics play a crucial role in determining species distributions and coexistence. Exploring causes of variation in seedling dynamics can therefore provide key insights into the factors affecting these phenomena. We examined the relative importance of biotic neighborhood processes and habitat heterogeneity using survival data for 5,827 seedlings in 39 tree and shrub species over 2?years from an old-growth temperate forest in northeastern China. We found significant negative density-dependence effects on survival of tree seedlings, and limited effects of habitat heterogeneity (edaphic and topographic variables) on survival of shrub seedlings. The importance of negative density dependence on young tree seedling survival was replaced by habitat in tree seedlings ??4?years old. As expected, negative density dependence was more apparent in gravity-dispersed species compared to wind-dispersed and animal-dispersed species. Moreover, we found that a community compensatory trend existed for trees. Therefore, although negative density dependence was not as pervasive as in other forest communities, it is an important mechanism for the maintenance of community diversity in this temperate forest. We conclude that both negative density dependence and habitat heterogeneity drive seedling survival, but their relative importance varies with seedling age classes and species traits.     

Strong distance-dependent effects for a spatially aggregated tropical species

A fundamental aspect of the Janzen–Connell Hypothesis (JCH) is that distance- and density-dependent mortality reduce the local dominance of species and cause regular rather than random or aggregated spatial patterns. Despite this explicit linkage between process and pattern, very few studies have explored how JCH processes translate into the spatial distributions of adult populations. In field experiments, we assessed germination, mortality and growth of conspecific and heterospecific seedlings beneath and away from Esenbeckia leiocarpa, a highly aggregated tropical tree species. We also investigated the effects of vertebrates using exclosures in the field, and the effects of pathogens in a soil sterilization shadehouse experiment. Germination of conspecifics underneath Esenbeckia was reduced by 64 % and mortality increased 28–123 % when compared to seedlings growing under other tree species; 99–100 % of Esenbeckia seedlings died under conspecifics. Heterospecifics were much less affected by Esenbeckia canopies. However, we found no evidence that either vertebrate herbivores or soil pathogens affected seed germination and seedling performance. Although many tropical tree species are aggregated, our results are the first to demonstrate strong negative distance-dependence for an aggregated species and one of the few to explore germination in the context of the JCH and show differences between conspecifics and heterospecifics, thus suggesting a broader role for Janzen–Connell processes as determinants of the distribution and abundance of tree species in tropical forests.     

Density dependence is not very prevalent in a heterogeneous subtropical forest

There is a growing body of evidence demonstrating that tree survival is influenced by negative density‐dependence, but it is still controversial how the effect may vary with life‐stage, and to what extent it plays a role in regulating tree survival in heterogeneous subtropical forests. In this study, we investigated density‐dependent effects on tree survival of six tree species in a 5‐ha subtropical forest in eastern China. The roughly 45 000 individuals in the forest were fully censused in 2003 and 2008. For each of these species, we used an inhomogeneous pair‐correlation function to quantify the change in spatial distribution for different size classes, and a case‐control design to study seedling–adult associations in 2003. Autologistic regression was used to determine the influence of neighborhood factors on individual survival from 2003 to 2008. We found that seedlings of five species were repulsed by distance to nearest conspecific adults in terms of their survival, consistent with predictions of the Janzen–Connell mechanism. By contrast, only the least shade‐tolerant Schima superba had a negative relationship with individual survival and conspecific distance‐weighted basal area. This suggests that the Janzen–Connell effect is only prevalent at the early seedling stage or seed‐to‐seedling phase. The strength of clustering significantly declined at sapling–pole and pole–adult transitions for Sycopsis sinensis and at seedling–sapling transition for Cleyera pachyphylla. Correlations between individual survival and conspecific abundance for these species were consistent with trends in the strength of clustering. These results suggest that density dependence plays a limited role in individual survival and species spatial structure beyond the early seedling stage (i.e. after true leaves growing) in this forest. In addition, this study indicates that including individuals from early life‐stages and factoring out potential confounding factors such as habitat preference are important in studies that seek evidence for density dependence in forest trees.     

Testing for Janzen–Connell Effects in a West African Montane Forest

The Janzen–Connell hypothesis proposes that density dependent seed and seedling mortality, combined with increasing seed and seedling survival away from the parent tree, together promote regular spacing of species and thus α diversity. This hypothesis has rarely been tested in tropical Africa, and rarely in montane forests anywhere. We tested this hypothesis using a combination of field experiments and observations in the most floristically diverse dry submontane forest in Nigeria. We investigated distance effects on seedling herbivory, seedling survival and seedling height growth. We found a significant decrease in herbivory with distance from conspecific adult trees for all three species of experimentally planted seedlings (Entandrophragma angolense, Deinbollia pinnata and Sterculia setigera), and also for naturally occurring seedlings of Pouteria altissima but not of Newtonia buchananii or Isolona pleurocarpa. The relative density of large seedlings/saplings of P. altissima, N. buchananii and I. pleurocarpa increased significantly at greater distance from conspecific adult trees; however, we found no significant distance effect on survival or height growth over 3 mo for all three experimentally planted species. Taken together, our results are some of the first to show that Janzen–Connell effects occur on the African continent and in particular montane tropical forest and suggest that such effects may be pantropical.     

Dispersion pattern and mortality of seeds and seedlings ofFagus crenata Blume in a cool temperate forest in western Japan

The seed and seedling mortality ofFagus crenata Blume after a mast year (1993) was examined in relation to density and distance from the nearest conspecific adult tree in a mixed conifer-hardwood forest in Ohdaigahara, western Japan. The mortality of fallen seeds during winter amounted to 93.7%, and 79.2% of the current-year seedlings died in the first growing season. The most important factor of death for both seeds and seedlings was predation by vertebrates. The mortality of seeds during winter was positively correlated with sound seed density. The mortality of seedlings was positively correlated with density but not significantly related to the distance from the nearest crown edge of a conspecific adult tree. Mortality patterns varied with stages and spatial scales due to the behavior of predators; it is thus important to investigate the spatial pattern of seeds and seedling mortality at various temporal and spatial scales. After the first growing season, the difference in seedling density between distance classes was not significant at <4m from the nearest adult trees due to density-dependent mortality. However, seedling density was significantly lower in the ≥4 m class than in the <4 m classes.     

Seedling survival in Manilkara butugi Chiov. along a dispersal gradient in Kakamega forest,Kenya

The spatial patterns of seedling distribution in Manilkara butugi were studied between December 2004 and April 2005 at Kakamega forest, Kenya. Our objectives were to determine the spatial variation in seedling density in relation to distance from the parent tree and the survivorship patterns of M. butugi seedlings. We hypothesized that seedling density and mortality would be highest beneath the parent tree and would reduce with increasing distance from the parent tree. Line transects, radiating along the four compass directions from mature female M. butugi trees were established, and quadrats marked out at intervals of 5 m in which seedlings were counted. To determine establishment and survivorship, belt transects 5 m × 40 m running along the northerly compass direction were laid and data collected on seedling variables. Seedling density reduced with increasing distance from the parent plant (F = 25.72, df = 3; P < 0.001). Whereas seedling height (F = 14.806, df = 1; P < 0.05) and the number of leaves (F = 12.45, df = 1; P < 0.05) increased with increasing distance from the parent tree, damage to seedlings reduced with increasing distance (F = 27.59, P < 0.05). Our results were consistent with the Janzen–Cornell escape hypothesis and have implications for natural regeneration of M. butugi and to maintenance of species diversity.     

Leaf diseases drive the Janzen–Connell mechanism regardless of light conditions: a 3-year field study

In forests, negative density/distance-dependent seedling mortality (NDD) caused by natural enemies plays a key role in maintaining species diversity [Janzen–Connell (J–C) model]. However, the relative importance of natural enemies in mediating NDD under heterogeneous light conditions has remained unclear. We examined the relative importance of pathogens (i.e., soil pathogens, leaf diseases) on seedling performance in forest understories (FUs) and gaps (gaps) during a 3-year period (results of first year of our study have been previously reported). For the hardwood, Prunus grayana, we investigated seedling mortality, morbidity agents, growth, and root infection by arbuscular mycorrhizal fungi (AMF) beneath conspecific and heterospecific adults in FUs and gaps. Seedling mortality was higher beneath conspecific than heterospecific adults throughout 3 years at both sites, mainly due to continuous leaf disease (i.e., angular leaf spot), whereas damping-off diseases caused mortality only in the first year. Beneath each adult, seedling mortality was higher in FUs than in gaps until second year, but it did not differ between two habitat types in the third year, because leaf diseases caused severe damage even in gaps. Seedling mass was significantly lower beneath conspecific adults. AMF infection of seedlings was also lower beneath conspecific adults, while it was higher in gaps than in FUs beneath both adults. This study demonstrates that the J–C model in a hardwood tree, P. grayana is mainly driven by high NDD seedling mortality caused by airborne leaf diseases, which continuously attack seedlings in a NDD manner regardless of environmental light conditions.     

Pathogen mortality of tropical tree seedlings: experimental studies of the effects of dispersal distance,seedling density,and light conditions

Summary We present results of two experiments designed to identify the relative importance of dispersal distance, seedling density, and light conditions on pathogen-caused mortality of tropical tree seedlings. The field experiment on Barro Colorado Island, Panama, demonstrated that both an increase in dispersal distance and a decrease in seedling density reduce levels of damping-off disease among seedlings of Platypodium elegans, and that there is an interaction between the two factors. The results indicated significant variation among sites in pathogen activity and suggested that seedlings are more vulnerable to disease when establishing around their parent tree than around other conspecific trees.The second experiment in a screened enclosure used potted seedlings of 18 wind-dispersed tree species exposed to two levels of sunlight and seedling density. The results indicated that environmental conditions similar to those in light-gaps significantly reduce pathogen activity. They also confirmed that high seedling density increases disease levels, especially under shaded conditions.Seedlings of 16 of the 18 species experienced pathogencaused mortality, but in widely varying amounts. Seed weight was not a good predictor of a species' vulnerability to pathogens. Adult wood density, an indicator of growth rate and successional status, was inversely correlated with a species' vulnerability to pathogens. Fast-growing, colonizing species, whose seedlings require light-gaps, lacked strong resistance to seedling pathogens, relative to slow-growing species able to tolerate shade and escape seedling pathogens. We discuss these results in the context of seed dispersal as a means of escaping from seedling pathogens.     

Short feeding period of carrot psyllid (Trioza apicalis) females at early growth stages of carrot reduces yield and causes leaf discolouration

Overwintered adult carrot psyllids [Trioza apicalis Förster (Homoptera: Psylloidea: Triozidae)] damage carrot [(Daucus carota ssp. sativum L.) (Apiaceae)] seedlings by phloem feeding on the leaves. The aim of this study was to investigate the carrot root and shoot growth in relation to carrot psyllid density during early growth stages. One, two, or three carrot psyllids were allowed to feed on carrot seedlings for 3 days. Leaf damage was measured at the 8‐leaf stage, and root, leaf fresh weight, and number of true leaves were measured at harvest. Both the age of the carrot seedling at infestation and the psyllid density had a significant effect on leaf damage at the 8‐leaf stage: seedlings damaged at the cotyledon stage exhibited more leaf damage than seedlings damaged at the 1‐leaf stage. A higher psyllid density significantly reduced the carrot root weight at harvest. The significant interaction of psyllid density with seedling age indicates that differently aged carrot seedlings responded differently to feeding: one psyllid feeding for 3 days at the cotyledon stage caused a significant yield loss, whereas three psyllids were needed to cause the same impact at the 1‐leaf stage. Carrot leaf weight at harvest was not reduced by carrot psyllid feeding: leaves recovered from the damage but roots did not. Our results confirm the farmers’ observations that a trap replacement period of 1 week for carrot psyllid monitoring is too long, especially at the cotyledon stage. Severe leaf discolouration on damaged carrots was observed at harvest. The possible reasons for this discolouration, such as toxin excreted in psyllid saliva or plant pathogenic mycoplasma infection, are discussed.     

Demography of the seedling bank of Manilkara zapota (L.) Royen,in a subtropical rain forest of Mexico

Selection of individuals in tropical trees, occurs mainly in the seedling phase, which in part explains the low densities of most species. The main objective of this work was to gain an insight into those factors that influence growth and survival of Manilkara zapota seedlings, one of the most abundant species in the lowland forests of Mexico and Central America. Eight 10-m² rectangular plots (5m×2m) were established, in which all < 35-cm-high M. zapota seedlings were marked, measured and enumerated at bimonthly intervals. In each census seedling height, number of leaves, length of largest leaf, type of damage and seedling death were registered. Seedling survival during 2 yr was high, reaching 82% including newly emerged seedlings. A maximum likelihood regression analysis showed that both number of leaves and length of largest leaf had a direct influence on seedling survival; however, neither type of damage nor seedling density and height had significant effect. Seedling height growth averaged 2.8 cm in the 2 yr of study. The combined effect of high seedling survival, reduced growth, and impact of physical damage mainly due to falling branches and leaves reveals the occurrence of a persistent seedling bank. Such a bank would contribute to recruitment of individuals in the juvenile and eventually in the adult stages. This seedling bank could explain the high density of individuals of M. zapota in the tropical forests of Mexico.     

Performance and fate of tree seedlings on and around nests of the leaf‐cutting ant Atta cephalotes: Ecological filters in a fragmented forest

Habitat fragmentation is currently the most pervasive anthropogenic disturbance in tropical forests and some species of leaf‐cutting ants of the genus Atta (dominant herbivores in the neotropics) have become hyper‐abundant in forest edges where their nests directly impact up to 6% of the forest area. Yet, their impacts on the regeneration dynamics of fragmented forests remain poorly investigated. Here we examine the potential of Atta cephalotes nests to function as ecological filters impacting tree recruitment. Growth, survival and biomass partitioning of experimentally planted seedlings (six tree species) were examined at eight spatially independent A. cephalotes colonies in a large Atlantic Forest fragment. Seedling performance and fate (leaf numbers and damage) were monitored up to 27 months across three habitats (nest centre, nest edge and forest understorey). Plants at illuminated nest centres showed twice the gross leaf gain as understorey individuals. Simultaneously, seedlings of all species lost many more leaves at nests than in the forest understorey, causing a negative net leaf gain. Net leaf gain in the shaded understorey ranged from zero (Licania and Thyrsodium species) to substantial growth for Copaifera and Virola, and intermediate levels little above zero for Protium and Pouteria. Also seedling survival differed across habitats and species, being typically low in the centre and at the edge of nests where seedlings were often completely defoliated by the ants. Lastly, seedling survival increased strongly with seed size at nest edges while there was no such correlation in the forest. Our results suggest that Atta nests operate as ecological filters by creating a specific disturbance regime that differs from other disturbances in tropical forests. Apparently, Atta nests favour large‐seeded tree species with resprouting abilities and the potential to profit from a moderate, nest‐mediated increase in light availability.     

Long-distance dispersal helps germinating mahogany seedlings escape defoliation by a specialist caterpillar

Herbivores and pathogens with acute host specificity may promote high tree diversity in tropical forests by causing distance- and density-dependent mortality of seedlings, but evidence is scarce. Although Lepidoptera larvae are the most abundant and host-specific guild of herbivores in these forests, their impact upon seedling distributions remains largely unknown. A firm test of the mechanism underpinning the Janzen–Connell hypothesis is difficult, even for a single tree species, because it requires more than just manipulating seeds and seedlings and recording their fates. Experimental tests require: (1) an insect herbivore that is identified and highly specialised, (2) linkage to an in situ measure (or prevention) of herbivory, and (3) evaluation and confirmation among many conspecific adult trees across years. Here we present experimental evidence for a spatially explicit interaction between newly germinating seedlings of a Neotropical emergent tree, big-leaf mahogany (Swietenia macrophylla, Meliaceae), and caterpillars of a noctuid moth (Steniscadia poliophaea). In the understory of a southeastern Amazon forest, the proportion of attacks, leaf area lost, and seedling mortality due to this specialised herbivore peaked near Swietenia trees, but declined significantly with increasing distance from mature fruiting trees, as predicted by the Janzen–Connell hypothesis. We conclude that long-distance dispersal events (>50 m) provided an early survival advantage for Swietenia seedlings, and propose that the role of larval Lepidoptera as Janzen–Connell vectors may be underappreciated in tropical forests.     

Comparison of Herbivores and Herbivory in the Canopy and Understory for Two Tropical Tree Species1

The Janzen–Connell model of tropical forest tree diversity predicts that seedlings and young trees growing close to conspecific adults should experience higher levels of damage and mortality from herbivorous insects, with the adult trees acting as either an attractant or source of the herbivores. Previous research in a seasonal forest showed that this pattern of distance‐dependent herbivory occurred in the early wet season during the peak of new leaf production. I hypothesized that distance‐dependent herbivory may occur at this time because the new foliage in the canopy attracts high numbers of herbivores that are limited to feeding on young leaves. As a consequence, seedlings and saplings growing close to these adults are more likely to be discovered and damaged by these herbivores. In the late wet season, when there is little leaf production in the canopy, leaf damage is spread more evenly throughout the forest and distance dependence disappears. I tested three predictions based on this hypothesis: (1) the same species of insect herbivores attack young and adult trees of a given plant species; (2) herbivore densities increase on adult trees during leaf production; and (3) herbivore densities in the understory rise during the course of the wet season. Censuses were conducted on adults and saplings of two tree species, raribea asterolepis and Alseis blackiana. Adults and saplings of both species had largely the same suite of chewing herbivore species. On adults of Q. asterolepis, the density of chewing herbivores increased 6–10 times during leaf production, but there was no increase in herbivore density on adults of A. blackiana. Herbivore densities increased 4.5 times on A. blackiana saplings and 8.9 times on Q. asterolepis saplings during the wet season, but there were no clear trends on the adults of either species. These results suggest that the potential of adult trees as a source of herbivores on saplings depends on the value of new leaves to a tree species' herbivores, which may differ across tree species.     

广西多穗柯叶片性状变异及幼苗生长量研究

多穗柯是一种珍贵天然野生药用植物,可以开发出保健食品色素和天然医用药品,广西的资源较丰富,该研究采集巴马、那坡、德保及田林等4个产地的多穗柯种子进行播种育苗,并跟踪调查测定一年生幼苗的叶片性状及幼苗生长量,并进行相关性分析。结果表明:(1)不同产地间叶片性状及幼苗生长指标均存在不同程度的差异,其中巴马与那坡、德保、田林在叶长、叶宽、叶面积、叶脉间距、叶鲜重、叶片干物质含量、叶片组织密度等叶片性状上的差异均达到显著水平,在株高、地径、单株干重、主根长、单株根干重及单株叶干重等生长指标上亦存在显著差异,且生长量是后3个产地的1~2倍;通过比较各产地的叶片保水力及植株净生长量,巴马的多穗柯植株耐旱性及生长速度优于其他三地。综合各性状表现,认为巴马的多穗柯苗期表现比较好,生长速度快,长势好,抗旱性较强,可作为多穗柯优良种源的初步选择。(2)8月份是多穗柯株高、地径的生长高峰期,建议此时应加强肥水管理,调节适宜的水肥光热条件,尽量延长幼苗的快速生长时间,以获得苗木的最大累积生长量。(3)叶片性状与幼苗生长量的相关性分析结果显示,叶面积与株高、地径、单株干重、单株根干重以及单株叶干重等呈极显著正相关,叶脉间距、叶绿素相对含量(SPAD)与株高、单株干重呈显著或极显著正相关,比叶面积与株高、地径呈显著负相关。因此,在以后的优株表型选择中,要优先考虑叶子大、叶脉间距宽、中老熟叶片叶色浓绿的植株。该研究结果为多穗柯优良种质资源的早期筛选提供了一定的依据。     

濒危植物闽楠幼苗更新及环境解释

为了更好地理解濒危植物闽楠幼苗天然更新与微环境的关系,以广西富川县蚌贝村闽楠天然林胸径5.0 cm以下的更新个体为研究对象。基于0.64 hm₂^{固定监测样地调查数据,采用Pearson相关系数、冗余分析等方法对幼苗更新密度与样地微环境因子关系进行分析。结果表明：(1)闽楠小幼苗更新个体数最多,随龄级增加更新密度呈下降趋势;幼苗向幼树转化率低,小幼苗向中幼苗转化率是闽楠更新成功的关键阶段;(2)闽楠幼苗更新密度与样地灌木层高度、最近母树距离、灌木层覆盖度极显著正相关,与样地光照、草本层覆盖度显著负相关; (3)灌木层高度、最近母树的距离、灌木层覆盖度、叶面积指数、光照、草本层覆盖度、凋落物厚度、土壤盐分是影响闽楠幼苗更新的主要微环境因子,光照和草本层覆盖度是闽楠幼苗更新的限制因子。因此,降低光照强度、减少草本层的覆盖度、增大灌木层的覆盖度等有利于闽楠种群更新发展。}     

Intraspecific and phylogenetic density-dependent seedling recruitment in a subtropical evergreen forest

Recent evidence suggests that plant performance can be influenced by the phylogenetic diversity of neighboring plants. However, no study to date has examined the effect of such phylogenetic density dependence on the transition from seed to seedling. Using 6 years of data on seedling recruitment and seed rain of 13 species from 130 stations (one 0.5 m² seed trap and three adjacent 1 m² seedling plots) in a subtropical evergreen forest, we asked: (1) Does negative density dependence act across seed to seedling stages? (2) Is there evidence for phylogenetic density dependence during the seed to seedling transition? (3) Does the strength of density dependence vary among years? Generalized linear mixed-effects models were used to model seed to seedling transition as a function of conspecific seed and seedling densities, heterospecific seed and seedling densities, and mean phylogenetic distance of heterospecific seeds and seedling. Conspecific seed density had a significant negative effect on seedling transition rates for 12 of 13 focal species. In contrast, conspecific seedling density had a positive effect for 7 species, suggesting species-specific habitat preferences. Few species were significantly affected by the density or phylogenetic relatedness of heterospecific seeds and seedlings. Only conspecific seed density effects varied among years for most focal species. Overall, our results reveal that conspecific seed and seedling densities play a more important role than the density or relatedness of heterospecific seeds and seedlings during the seed to seedling stage, suggesting that species-specific seed predators, along with habitat preferences, may contribute to diversity maintenance in this forest.     

Invertebrate predators drive distance-dependent patterns of seedling mortality in a temperate tree Acer pseudoplatanus

There is growing evidence that distance- and density-dependent recruitment are important factors in structuring tree communities in temperate forests. Although pathogens have generally been assumed to be driving these patterns, few studies have attempted to experimentally identify the agent responsible. Here we investigate patterns of recruitment in sycamore Acer pseudoplatanus , a temperate tree species, in its exotic range in Britain. Seedling mortality, the limiting stage in sycamore regeneration, was monitored with respect to the proximity and density of adult sycamores and seedlings in a grid of sampling units. We found that mortality was significantly greater beneath adult sycamores compared with heterospecifics, and was further elevated in areas of high conspecific basal area, a proxy for biomass. An experimental approach that manipulated the access of predators to seedlings demonstrated that herbivorous invertebrates were responsible for driving these distance- and biomass-dependent trends in mortality. Evidence of stem damage on dead seedlings suggests that slugs are the principal cause of mortality, although there was only partial evidence of higher slug activity beneath sycamores. The implications of these findings for sycamore invasion, species co-existence and management are discussed.     

Tree seedling richness,but not neighborhood composition,influences insect herbivory in a temperate deciduous forest community

Insect herbivores can serve as important regulators of plant dynamics, but their impacts in temperate forest understories have received minimal attention at local scales. Here, we test several related hypotheses about the influence of plant neighborhood composition on insect leaf damage in southwestern Pennsylvania, USA. Using data on seedlings and adult trees sampled at 36 sites over an approximately 900 ha area, we tested for the effects of total plant density, rarefied species richness (i.e., resource concentration and dietary‐mixing hypotheses), conspecific density (i.e., Janzen–Connell hypothesis), and heterospecific density (i.e., herd‐immunity hypothesis), on the proportion of leaf tissue removed from 290 seedlings of 20 species. We also tested for the effects of generic‐ and familial‐level neighborhoods. Our results showed that the proportion of leaf tissue removed ranged from zero to just under 50% across individuals, but was generally quite low (<2%). Using linear mixed models, we found a significant negative relationship between insect damage and rarefied species richness, but no relationship with neighborhood density or composition. In addition, leaf damage had no significant effect on subsequent seedling growth or survival, likely due to the low levels of damage experienced by most individuals. Our results provide some support for the resource concentration hypothesis, but suggest a limited role for insect herbivores in driving local‐scale seedling dynamics in temperate forest understories.     

Distance- and density-dependent seedling mortality caused by several diseases in eight tree species co-occurring in a temperate forest

To examine whether the Janzen–Connell mechanism applies to temperate forests, seedling survival and causes of mortality were investigated at two distances (beneath, far) from conspecific adults and at two densities (high, low) at each distance for seedlings (n = 7935) of eight tree species co-occurring in a hardwood forest. Six of the eight species showed distance- and/or density-dependent seedling mortality mainly caused by diseases and rodents. In four of the five species primarily killed by disease (i.e. damping-off, blight, rot, powdery mildew), the infectivity (probability of infection by the disease) and/or the virulence (proportion of seedlings killed to those infected by the disease) were higher beneath than far from conspecific adults. These findings suggest that host specificity and/or spatially heterogeneous activity of natural enemies play an important role in the reciprocal replacement of tree species, maintaining species diversity in temperate forests.     

Distribution ofAcer palmatum seedlings under the crown of the maternal tree

To investigate the relationship between crown width of a parent plant and seedling distribution, seedling ages and sizes, and their distances from the maternal tree were surveyed under a maple (Acer palmatum) crown at Chungdam Park, Chungdam-dong, Kangnam-gu, Seoul, in 1998. Maple crown width increased 33.0 cm per year during the eight-year study. Seedlings ranged from 1 to 10 years old; their density was highest the third year, and decreased with age. Seedling-distribution ranges from the parent plant were 43 cm to 487 cm, and seedling density was highest at the border of the parent plant canopy. One-year-old seedlings were distributed broadly, but the main distribution areas became more distant from the maternal plant with seedling age. Seedling heights increased with age; differences between two consecutive ages were significant at the 0.1% level for ages less than 5 years, but not for over 6 years. For seedlings of the same age, heights were similar, but short under the tree crown, but were varied and taller outside the crown. The variation in seedling height increased with age and distance from the parent plant In particular, seedlings older than 5 years that were distributed outside the crown showed conspicuous variations among distance classes. Therefore, maple seedlings were distributed to an appropriate distance at which their growth could be supported by the understory-light environment. This study demonstrated that newA. palmatum trees are not recruited from the seedlings growing beneath the crown but from those that grow vigorously outside the crown.