Conspecific Seedlings Research Articles

Intense light and conspecific density increase seedling mortality across age groups in a typhoon‐disturbed tropical forest

AbstractQuestionsMaintaining forest species diversity, regenerative function and ecosystem resilience begins with successful seedling establishment. Despite their critical role, seedlings face the highest mortality risks among plant life‐history stages. Our research aims to answer the following questions: In a tropical forest disturbed by typhoons, what is the median survival time for seedlings, to what extent do light levels contribute to mortality risks and what are the crucial mortality risks across seedling age groups?LocationA tropical mountain zonal foothill evergreen broad‐leaved forest in Nanjenshan Nature Reserve, Taiwan, frequently visited by typhoons.MethodsWe investigated woody seedlings in 75 quadrats (each 1 m × 1 m) every 3 months from October 2009 to January 2018, spanning 8.3 years with 34 censuses. We used the Kaplan–Meier method to estimate seedlings’ median survival time and Generalized Linear Models to identify mortality risk factors for all seedlings and those that persisted beyond the median survival time.ResultsThe median survival time for all seedlings was 0.5 years, ranging from 0.25 years for shade‐intolerant species to 1.25 years for shade‐tolerant species. Seedlings germinated in high‐light microhabitats and high conspecific seedling density exhibited higher mortality probabilities, regardless of age.ConclusionsThis study reveals the short survival time and key mortality risks of seedlings in a typhoon‐disturbed forest. The negative impact of light levels on seedling survival highlights the potential consequences of increased light levels resulting from typhoon disturbances in the study region.

Drought tolerance and species abundance mediate dry season negative density dependence in a tropical forest.

Conspecific negative density dependence (CNDD) is thought to be a key process in maintaining plant diversity. However, the strength of CNDD is highly variable in space and time as well as among species, and correlates of this variation that might help to understand and explain it remain largely unquantified. Using Bayesian hierarchical models, we took advantage of 10-year seedling monitoring data that were collected annually in every dry and rainy season in a seasonal tropical forest. We quantified the interspecific variation in the strength of CNDD and its temporal variation. We also examined potential correlates of this interspecific and temporal variation, including species functional traits (such as drought-tolerant traits, defense-related traits, and recourse acquisition traits) and species abundances. In the dry season, we found a negative relationship between the density of neighboring conspecific seedlings on seedling survival, while in the rainy season, there was a negative relationship between the density of neighboring conspecific adults on seedling survival. In addition, we found that interspecific variation in CNDD was related to drought-tolerant traits in the dry season but not in the rainy season. Across years, we found that drought-intolerant species suffer less CNDD during the dry seasons that have higher rainfall, whereas drought-tolerant species suffer less CNDD when the dry season has lower rainfall. We also found that rare species suffered stronger CNDD in the dry season. Overall, our study highlights that CNDD is highly variable among species and through time, necessitating a deeper appreciation of the environmental and functional contexts of CNDD and their interactions.

Plant-ant interactions mediate herbivore-induced conspecific negative density dependence in a subtropical forest

The early growth stage of plants is vital to community diversity and community regeneration. The Janzen-Connell hypothesis predicts that conspecific density dependence lowers the survival of conspecific seedlings by attracting specialist natural enemies, promoting the recruitment and performance of heterospecific neighbors. Recent work has underscored how this conspecific negative density dependence may be mediated by mutualists – such as how mycorrhizal fungi may mediate the accrual of host-specific pathogens beneath the crown of conspecific adult trees. Aboveground mutualist and enemy interactions exist as well, however, and may provide useful insight into density dependence that are as of yet unexplored. Using a long-term seedling demographic dataset in a subtropical forest plot in central China, we confirmed that conspecific neighborhoods had a significant negative effect on seedling survival in this subtropical forest. Furthermore, although we detected more leaf damage in species that were closely related to ants, we found that the presence of ants had significant positive effects on seedling survival. Beside this, we also found a negative effect of ant appearance on seedling growth which may reflect a trade-off between survival and growth. Overall, our findings suggested that ants and conspecific neighborhoods played important but inverse roles on seedling survival and growth. Our results suggest ants may mediate the influence of conspecific negative density dependence on seedling survival at community level.

Foliar disease incidence in a tropical seedling community is density dependent and varies along a regional precipitation gradient

Abstract Many studies identify fungal and oomycete phytopathogens as natural enemies capable of influencing plant species composition and promoting diversity in plant communities. However, little is known about how plant‐pathogen interactions vary along regional abiotic gradients or with tree species characteristics, which limits our understanding of the causes of variation in tree species richness. We surveyed 10,756 seedlings from 272 tree species for disease symptoms along a mean annual precipitation gradient in the tropical wet forests of Central Panama for 3 months in the early wet season (June–August) and 2 months in the following dry season (March–April). Over 99% of observed disease symptoms were caused by necrotrophic foliar pathogens, while less than 1% of symptoms were attributed to soilborne pathogens. Foliar disease incidence was inversely related to mean annual precipitation, a pattern which may be due to greater disease susceptibility among dry forest species. Foliar disease incidence increased with conspecific seedling density but did not respond to the proximity of conspecific adults. Although foliar disease incidence decreased as mean annual precipitation increased, the strength of conspecific density‐ or distance‐dependence was independent of the precipitation gradient. Seedlings of common tree species and species dispersed by non‐flying mammals had a higher risk of foliar pathogen incidence. Increased disease in common species may help reduce their dominance. Synthesis. The increases in foliar pathogen incidence with conspecific seedling density, species abundance, and dispersal mechanism indicate that foliar disease incidence is non‐random and may contribute to the regulation of tropical plant communities and species coexistence. Furthermore, the relationships between foliar disease incidence, dispersal mechanism and precipitation suggest plant‐pathogen interactions could shift as a response to climate change and disruption of the disperser community.

Travelling waves due to negative plant–soil feedbacks in a model including tree life-stages

The emergence and maintenance of tree species diversity in tropical forests is commonly attributed to the Janzen–Connell (JC) hypothesis, which states that growth of seedlings is suppressed in the proximity of conspecific adult trees. As a result, a JC distribution due to a density-dependent negative feedback emerges in the form of a (transient) pattern where conspecific seedling density is highest at intermediate distances away from parent trees. Several studies suggest that the required density-dependent feedbacks behind this pattern could result from interactions between trees and soil-borne pathogens. However, negative plant–soil feedback may involve additional mechanisms, including the accumulation of autotoxic compounds generated through tree litter decomposition. An essential task therefore consists in constructing mathematical models incorporating both effects showing the ability to support the emergence of JC distributions.In this work, we develop and analyse a novel reaction–diffusion-ODE model, describing the interactions within tropical tree species across different life stages (seeds, seedlings, and adults) as driven by negative plant–soil feedback. In particular, we show that under strong negative plant–soil feedback travelling wave solutions exist, creating transient distributions of adult trees and seedlings that are in agreement with the Janzen–Connell hypothesis. Moreover, we show that these travelling wave solutions are pulled fronts and a robust feature as they occur over a broad parameter range. Finally, we calculate their linear spreading speed and show its (in)dependence on relevant nondimensional parameters.

Seedling performance in a dioecious tree species is similar near female and male conspecific adults despite differences in colonization by arbuscular mycorrhizal fungi

Plant–soil feedbacks (PSFs) are a key driver of species diversity and composition in plant communities worldwide; however, the factors that may cause feedbacks to vary within species are rarely examined. In dioecious species, the strength of feedbacks may differ near female plants that produce seed versus near male plants (which do not) because repeated inputs of seeds and high seedling densities near females may cause accumulation of host‐specific soil microbes that influence seedling performance. To test whether conspecific seedling performance is reduced near seed‐producing female trees relative to male or heterospecific trees, we conducted shadehouse and field experiments with a dioecious tropical tree species, Virola surinamensis (Myristicaceae), on Barro Colorado Island, Panama. The shadehouse experiment isolated the effect of soil microbial communities on seedling growth and allowed us to quantify colonization by mutualistic arbuscular mycorrhizal (AM) fungi, while the field experiment allowed us to assess seedling survival and growth in the presence of nearby conspecific adults and seedlings. In both experiments, seedling performance was similar between seedlings grown in the soil microbial communities and field environments underneath female conspecific, male conspecific, and heterospecific trees. However, contrary to expectation, seedling colonization by AM fungi was higher in male conspecific soil microbial communities than in female or heterospecific soil microbial communities at the end of the shadehouse experiment. Together, our experiments show that while differences among female and male plants in dioecious species may influence the association of conspecific seedlings with AM fungi in their soils, this variation does not necessarily translate directly to differences in seedling performance, at least over the time frame of our experiments. Studies of additional dioecious species are needed to help determine differences in soil microbial communities beneath male and female plants and to assess the role of seed input versus adult root systems in driving PSFs.

Effects of tree species diversity and conspecific seedling density on insect herbivory and pathogen infection on big‐leaf mahogany seedlings

The Janzen–Connell hypothesis (JCH) predicts that attack by specialist enemies on seedlings increases with conspecific seedling density, but studies have rarely experimentally tested for the contingency of such effects on the tree community context (e.g. diversity, composition) ormeasured responses by different enemies (e.g. herbivores, pathogens). We conducted a field study in a large‐scale system evaluating tree species diversity and conspecific density effects on leaf damage on mahogany Swietenia macrophylla seedlings. We established quadrats of eight levels of seedling density across mahogany tree monocultures and tree species polycultures including mahogany, and recorded percent leaf damage by insects and percent leaf necrosis by a pathogenic fungus on mahogany seedlings. We found contrasting effects of tree species diversity on insects and pathogens. Whereas tree diversity did not affect leaf damage by insects, it had a significant negative effect on leaf necrosis by pathogens whereby, on average, percent leaf necrosis on mahogany seedlings in polyculture was half of that observed in monoculture. We discuss the potential influences of changes in mahogany tree density versus frequency (relative to other tree species) driving this diversity effect. Unexpectedly, we found no effect of seedling conspecific density on either leaf insect or pathogen damage (i.e. density‐independent attack). Likewise, we found no significant tree diversity by seedling density interaction, indicating that plant enemies were consistently unresponsive to variation in seedling density acrosslevels of tree diversity. Overall, this study provides a unique test of the JCH by experimentally evaluating seedling density and tree diversity effects on contrasting plant enemies and, in turn, the controls they exert over plant recruitment.

Effects of tree density and herbaceous plants on tree seedling survival across the growing and non-growing season in a temperate forest

Tree seedling survival is critical for estimating forest recruitment, but it remain elusive about how neighboring tree’s density and herbaceous plants affect it. To assess the relative importance of tree density and herbaceous plants on tree seedling survival during the growing season and non-growing season, we established a two-year experiment with seedling census and removal of herbaceous plants in both secondary and old-growth temperate forest sites in the temperate forests of northeastern China. A total of 3,443 seedlings in 17 tree species were surveyed in this study. We found that both tree density and herbaceous plants significantly affected tree seedling survival, but the specific effects depended on shade tolerance, seedling growth stage and growing season. Conspecific adult neighbors had significant negative effects on the survival of shade-tolerant seedlings during the growing season, while conspecific seedling densities positively correlated with shade-intolerant seedlings. Herb removal significantly improved the survival of newly germinated and shade-intolerant seedlings during the growing season. Furthermore, we found that the survivals of both newly germinated and established seedlings differed seasonally in response to adult, seedling neighbors and herb removal. Herb removal enhances seedling survival during the non-growing season by offsetting the negative effects of conspecific adult neighbors during the growing season. Our findings provide new evidence for seedlings’ shade tolerance, growth stage and seasonal dependence of density dependence in temperate forests, and highlight the significance of herbaceous plants for seedling survival, an important factor that has been neglected in many previous studies.

Roles of pathogens and mycorrhizae in conspecific negative distance dependency and replacement of tree species in a temperate forest

Maintaining and recovering species diversity is one of the main management goals of forest managers. It is thus important to understand the mechanisms that drive species diversity. The mechanism is strongly related to the harsh biological environment surrounding the tree, especially its interaction with natural enemies, and a well-known mechanism is the Janzen-Connell hypothesis. The hypothesis is best understood by evaluating not only conspecific negative distance-dependent (CNDD) seedling mortality but also replacement of conspecific seedlings by heterospecifics. However, it remains unclear how antagonistic pathogens and mutualistic mycorrhizae affect the strength of replacement. A reciprocal seed-sowing experiment was conducted for five tree species with different mycorrhizal types in forest understorey (FU) and gaps (Gap) in a temperate forest. We investigated seedling mass, mortality, mortality agents, and root infection with arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) fungi. In both FU and Gap sites, two AM species were attacked by pathogens causing damping-off and leaf diseases more severely beneath conspecific than heterospecific adults, and more severely among conspecifics than heterospecifics of focal adults, resulting in strong CNDD and replacement, whereas seedling mortality in two EM species due to damping-off pathogens and leaf diseases did not differ among treatments, resulting in weak CNDD and replacement. One AM species showed an intermediate trend between two AM species and two EM species. Thus, CNDD and replacement caused by pathogens were positively correlated in FU but not Gap sites. These traits reveal that replacement closely correlated with CNDD play an important role in maintaining species diversity, especially when gaps are absent. This study also suggest that replacement plays an important role in regulating the spatial distribution of individual trees and subsequent relative abundance of tree species, as well as CNDD effects. Forest managers need to consider differences in the strength of CNDD and replacement between mycorrhizal fungi types when attempting to achieve successful seedling establishment after harvesting without compromising species diversity.

Distance and density dependence in two native Bornean dipterocarp species.

The Janzen-Connell hypothesis proposes that density and distance-dependent mortality generated by specialist natural enemies prevent competitive dominance. Much literature on Janzen-Connell mechanisms comes from the neotropics, and evidence of the role of distance and density-dependence is still relatively sparse. We tested the predictions of the Janzen-Connell hypothesis in a South-East Asian system dominated by mast fruiting species. We hypothesized that seedling survival would decrease with distance and density, seedling growth would increase, and herbivory would decrease, according to the predictions of the Janzen-Connell hypothesis. Experiments were conducted to determine the strength of the Janzen-Connell mechanism by manipulating the density and identity of tree species as a function of the distance from parent trees. Survival of conspecific seedlings was reduced near adult trees of one species, but not another. High densities of seedlings decreased the growth of conspecific seedlings of both species. In both species, herbivory rates decreased with distance in low-density areas. This study indicates that dipterocarp species experienced weak Janzen-Connell effects of distance and density dependence at the growth stage studied. Future studies in this system might focus on earlier life-history stages such as seeds and small seedlings, as well as studying mortality during mast-seeding events.

Legacy effects post removal of a range-expanding shrub influence soil fungal communities and create negative plant-soil feedbacks for conspecific seedlings

Legacy effects post removal of a range-expanding shrub influence soil fungal communities and create negative plant-soil feedbacks for conspecific seedlings

Tree mycorrhizal type mediates conspecific negative density dependence effects on seedling herbivory, growth, and survival.

Tree mycorrhizal type plays an important role in promoting plant species diversity and coexistence, via its mediating role in conspecific negative density dependence (CNDD), i.e., the process by which an individual's performance is impaired by the density of conspecific plants. Previous findings suggest that ectomycorrhizal (EM) tree species are generally less susceptible to CNDD than arbuscular mycorrhizal (AM) tree species, due to the chemical and physical protection that EM fungi provide their host with. We examined how CNDD effects on leaf herbivory, seedling growth, and survival differ between AM and EM seedlings of ten tree species collected over 3years in an old-growth temperate forest in northeastern China. We found that AM and EM seedlings differed in how conspecific density affected their leaf herbivory, seedling growth, and survival. Specifically, AM seedlings leaf herbivory rates significantly increased with increasing conspecific seedling and adult density, and their growth and survival rates decreased with increasing conspecific adult density, these patterns were, however, absent in EM seedlings. Our work suggests that AM seedlings have a performance disadvantage relative to EM seedlings related to the negative effects from conspecific neighbors. We highlight the importance of integrating information on seedling leaf herbivory, seedling growth, to provide further understanding on potential mechanisms driving differences in CNDD between AM and EM tree seedlings.

Arbuscular mycorrhizal fungi improve legume seedling performance as the competition with conspecific neighbouring adult plants decreases

In temperate grasslands, the balance between competitive and facilitative interactions between adult plants and conspecific seedlings can be affected by defoliation intensity, soil nutrient deficiency and arbuscular mycorrhizal fungi (AMF). Here, we investigated the effect of AMF on the interactions between Lotus tenuis Waldst. & Kit. seedlings grown in the presence or absence of conspecific adult plants defoliated at different intensities (0, 50 and 75%) in a P-deficient soil of the Salado River Basin (Argentina). Mycorrhizal seedlings grown near defoliated and non-defoliated adult plants produced less shoot dry weight (DW) than non-mycorrhizal counterparts, whereas mycorrhizal seedlings alone showed 258% and 504% more shoot DW and shoot P content, respectively than non-mycorrhizal counterparts. This was associated with higher growth and P uptake of mycorrhizal adult plants than non-mycorrhizal counterparts. The competition index between seedlings and adult plants showed competition interaction in mycorrhizal systems (from −2.6 to −1.9) and neutral interaction in non-mycorrhizal systems (near to zero) despite the defoliation intensity. We propose that moderate defoliation (50%) allows maintaining high mycorrhizal benefits in adult plants (110.1% and 168.8% mycorrhizal growth and P response, respectively), whereas high defoliation (75%) decreases mycorrhizal benefits (27.18% and 80.21% mycorrhizal growth and P response, respectively). We highlight AMF symbiosis as an important strategy of L. tenuis adult plants to regulate the growth of neighbouring conspecific seedlings in a P-deficient soil.

Wind dispersal and 1‐year survival of Vataireopsis iglesiasii (Fabaceae) seedlings in a Neotropical lowland rain forest

AbstractPopulations of many tropical tree species are regulated by negative distance‐ and density‐dependent processes (NDD), yet most studies on the effects of conspecific seedling and adult neighbors on seedling survival have focused on animal‐dispersed species. Species with seeds dispersed by wind may not be moved as far on average as seeds dispersed by animals, but some seeds may be dispersed a lot further, suggesting that knowledge of dispersal mechanism may help in our understanding of NDD. In this study, we took advantage of a high‐fecundity reproductive event that occurred for an individual isolated canopy tree of Vataireopsis iglesiasii in a tropical lowland rain forest site in Amazonian Ecuador to document seed dispersal and seedling survival to 1‐year post‐dispersal. Most seeds did not disperse far: 86% of germinated seedlings were found within 100 m. Mortality was high: only 49 of the 1732 monitored seedlings survived one year and only five survived a further four years. We found a significant negative effect of conspecific seedling density (but no effect of distance from the parent tree) on 1‐year survival. Rare long‐distance dispersal events may increase the probability of a seed reaching specific habitats, such as high‐light patches, and surviving beyond one year, thereby shifting the population recruitment curve outwards away from adult trees and maintaining diversity in species‐rich forests.Abstract in Spanish is available with online material.

Variation in biotic interactions mediates the effects of masting and rainfall fluctuations on seedling demography in a subtropical rainforest

Abstract Seedling dynamics are critical for determining community composition and future forest structure. Two mechanisms are crucial drivers of seedling vital rates: biotic interactions and environmental filtering. Biotic interactions at the neighbourhood scale, such as the Janzen–Connell effect, are considered among the main mechanisms maintaining species coexistence. These effects can show considerable temporal variation because of changes in neighbourhood density (i.e. seedling recruitment following masting years) and weather conditions. However, there is limited evidence of whether interannual variation in these conditions modulates biotic interactions and its potential cascading effects on seedling vital rates. We used seedling data collected over 17 years in a subtropical rainforest in northern Taiwan. We modelled interannual variation in first‐year survival and growth of 12,830 seedlings of 36 species and 47 cohorts using (generalized) linear mixed‐effects models with several crowding indices and abiotic conditions (edaphic and topographic factors) as predictors. The first‐year seedling survival and growth depended on both weather conditions and neighbourhood density. Seedling survival was greater when recruitment was more abundant and in years with greater rainfall, and this was mostly mediated by the interaction with conspecific and heterospecific seedlings. Seedling growth decreased with the increasing abundance of new recruits and increased with annual rainfall. Growth was negatively affected by the basal area of conspecific and heterospecific trees and by the density of conspecific seedlings when the number of new recruits was greater. The abiotic factors had a limited effect on seedling survival and growth compared to biotic factors. Synthesis. Our results show that variation in recruitment abundance due to masting and annual rainfall significantly affects seedling vital rates by changing biotic interactions. These findings are in accordance with the economy of scale hypothesis, which predicts higher per‐capita reproductive success in years of higher seed production. Furthermore, the results support the hypothesis that fluctuations in rainfall can alter biotic interactions in forests. Efforts to predict forest community responses to climate changes should consider the temporal variations in biotic interactions.

Oomycetes associated with Prunus serotina persist in soil after tree harvest

Soil-borne pathogens can shape forest communities by lowering seedling survivorship. Many soil pathogens can persist long-term as survival spores, but how long pathogens outlive tree hosts in gap soils and whether they continue to affect seedling survival is uncertain. We studied the presence of oomycetes and evaluated seedling performance in soils near live Prunus serotina trees, and 0.5 and 1.5-y-old stumps. We isolated five species of oomycetes from soils, two of which were pathogenic (Pythium intermedium and Pythium irregulare) to Prunus serotina. There was a non-significant ~10.5% increase in conspecific seedling survival in stumps versus live trees, and pathogens were present in soils of all stump ages. The continued presence of pathogens of Prunus serotina in gap soils demonstrates the potential for impacts on conspecific regeneration after tree death, though the slight improvement in survival suggests that these effects may weaken with time.

Short-lived legacies of Prunus serotina plant-soil feedbacks.

Plant-soil feedbacks (PSFs) are often involved in fundamental ecological processes such as plant succession and species coexistence. After a plant initiating PSFs dies, legacies of PSFs occurring as soil signatures that influence subsequent plants could persist for an unknown duration. Altered resource environments following plant death (especially light availability) could affect whether legacy effects manifest and persist. To evaluate PSFs and their legacies, we obtained soils from a chronosequence of Prunus serotina harvests. In a greenhouse experiment, we planted conspecific seedlings under two light levels in these soils of varying time since the influence of live Prunus serotina, and compared seed/seedling survival in soils from live trees, stumps, and surrounding forest matrix within each site and across the chronosequence. PSF legacies were measured as the difference between seedling performance in live tree and stump soils within a site. Negative PSF legacies of P. serotina were short-lived, lasting up to 0.5years after tree removal. These effects occurred under 5% but not 30% full sun. PSFs and their legacies manifested in seed/seedling survival, but not biomass. Though restricted to low light, short-lived legacies of P. serotina PSFs could have lasting impacts on plant community dynamics during post-disturbance regeneration by disfavoring P. serotina regeneration in small tree-fall gaps.

Environmental context alters the magnitude of conspecific negative density dependence in a temperate forest

AbstractConspecific negative density dependence (CNDD) is an important driver of stand‐level tree diversity in temperate forests. CNDD can occur via two main processes. First, soil pathogens and herbivores form a halo of high enemy density surrounding established trees at local spatial scales. As a result, seedlings and saplings residing within this halo experience reduced growth and survival. Second, intraspecific competition can lead to local depletion of resources, reducing the performance of conspecific seedlings and saplings, and encouraging spatial mixing of tree species. While seedlings and saplings respond to nearby conspecific adults, they are simultaneously responding to local environmental conditions. Solar radiation, soil moisture, and soil nutrients can all influence tree recruitment patterns. Furthermore, these environmental factors can influence densities and activities of soil pathogens, as well as seedling and sapling carbon balance. As a result, we expect small‐scale variation in the abiotic environment to influence the magnitude of CNDD, although this expectation has thus far received little attention. We combined tree stem and environmental data from a mapped oak‐hickory forest plot to evaluate whether the local environment modulates the direction or magnitude of negative density dependence. We tested for the interaction between conspecific and heterospecific basal area within a sapling's neighborhood and each of five environmental covariates: index of canopy gaps, soil organic matter, soil cations, topographic curvature, and topographic slope/aspect. We found that increasing local conspecific and heterospecific basal areas reduced sapling growth and that the conspecific basal area reduced sapling growth more so than did heterospecific basal area. Furthermore, we observed significant interactions between adult tree density and all environmental covariates that differed depending on the category of nearby adults (i.e., conspecific vs. heterospecific), and this effect was most important with variations in soil organic matter, soil cations, and topographic slope/aspect. We conclude that tree basal area not only interacts with environmental conditions to influence sapling growth, but more importantly, that small‐scale environmental conditions interact with the density of nearby conspecifics to modulate sapling growth. Thus, if this result proves general, a central mechanism underlying the maintenance of diversity can be modified by small‐scale environmental conditions.

The dynamics of conspecific tree and seedling neighbors on seedling survival in a subtropical forest

Negative intraspecific interactions could provide opportunities for heterospecific regeneration, thereby maintaining species coexistence in forest communities. Increasing conspecific tree and seedling neighbor densities often have negative correlations with seedling survival. If and how the strength of interactions change with seedling age remains a knowledge gap. In this study, we followed 2184 newly germinated seedlings belonging to 26 species for five years in a subtropical evergreen broad-leaved forest in eastern China. We tested the density distribution pattern of different aged live seedlings and used generalized linear mixed models to assess the dynamic influence of conspecific tree and seedling neighbors on seedlings survival across years. Our results showed that the density distribution of newly germinated seedlings was more clustered than the other aged live seedlings. Over five years, the clustered density distribution of live seedlings gradually weakened. Particularly, the density distribution of five years old seedlings was statistically random. Meanwhile, the degree of positive correlation between the density of live seedlings and the density of conspecific tree neighbors decreased with seedling age. Correspondingly, the conspecific seedling neighbors had a significant negative correlation with newly germinated seedlings survival in the first year and had a greater effect than conspecific tree neighbor density. As seedlings age, the conspecific tree neighbors had a greater negative relative effect on seedlings survival than the conspecific seedling neighbors. Our findings demonstrate that intraspecific negative interactions are an important mechanism on seedling survival and the magnitude of those interactions change with seedling age. In younger seedlings, intraspecific negative interactions were mainly from conspecific seedling neighbors and weakened as the seedlings aged. In later seedling ages, the interactions from conspecific tree neighbors dominated. Ultimately, these effects reduce species clumping dynamically and provide space for establishment of heterospecifics which would maintain diversity in the forest.

Limited contributions of plant pathogens to density-dependent seedling mortality of mast fruiting Bornean trees.

Fungal pathogens are implicated in driving tropical plant diversity by facilitating strong, negative density‐dependent mortality of conspecific seedlings (C‐NDD). Assessment of the role of fungal pathogens in mediating coexistence derives from relatively few tree species and predominantly the Neotropics, limiting our understanding of their role in maintaining hyper‐diversity in many tropical forests. A key question is whether fungal pathogen‐mediated C‐NDD seedling mortality is ubiquitous across diverse plant communities. Using a manipulative shadehouse experiment, we tested the role of fungal pathogens in mediating C‐NDD seedling mortality of eight mast fruiting Bornean trees, typical of the species‐rich forests of South East Asia. We demonstrate species‐specific responses of seedlings to fungicide and density treatments, generating weak negative density‐dependent mortality. Overall seedling mortality was low and likely insufficient to promote overall community diversity. Although conducted in the same way as previous studies, we find little evidence that fungal pathogens play a substantial role in determining patterns of seedling mortality in a SE Asian mast fruiting forest, questioning our understanding of how Janzen‐Connell mechanisms structure the plant communities of this globally important forest type.