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.     

Seedling survival simultaneously determined by conspecific,heterospecific, and phylogenetically related neighbors and habitat heterogeneity in a subtropical forest in Taiwan

Density dependence and habitat heterogeneity have been recognized as important driving mechanisms that shape the patterns of seedling survival and promote species coexistence in species‐rich forests. In this study, we evaluated the relative importance of density dependence by conspecific, heterospecific, and phylogenetically related neighbors and habitat heterogeneity on seedling survival in the Lienhuachih (LHC) Forest, a subtropical, evergreen forest in central Taiwan. Age‐specific effects of different variables were also studied. We monitored the fates of 1,642 newly recruited seedlings of woody plants within a 25‐ha Forest Dynamics Plot for 2 years. The effects of conspecific, heterospecific, and phylogenetically related neighbors and habitat heterogeneity on seedling survival were analyzed by generalized linear mixed models. Our results indicated that conspecific negative density dependence (CNDD) had a strong impact on seedling survival, and the effects of CNDD increased with seedling age. Heterospecific positive density dependence (HPDD) and phylogenetic positive density dependence (PPDD) had a significant influence on the survival of seedlings, and stronger HPDD and PPDD effects were detected for older seedlings. Furthermore, seedling survival differed among habitats significantly. Seedling survival was significantly higher in the plateau, high‐slope, and low‐slope habitats than in the valley. Overall, our results suggested that the effects of CNDD, HPDD, PPDD, and habitat heterogeneity influenced seedling survival simultaneously in the LHC subtropical forest, but their relative importance varied with seedling age. Such findings from our subtropical forest were slightly different from tropical forests, and these contrasting patterns may be attributed to differences in abiotic environments. These findings highlight the importance to incorporate phylogenetic relatedness, seedling age, and habitat heterogeneity when investigating the impacts of density dependence on seedling survival that may contribute to species coexistence in seedling communities.     

长白山次生针阔混交林乔木幼苗存活的影响因素分析

基于长白山次生针阔混交林样地, 以520个1 m × 1 m幼苗样方中胸径小于1 cm的乔木幼苗为研究对象, 选取2013年和2014年的幼苗调查数据, 运用广义线性混合模型(GLMM)分析了生物邻体和生境异质性对幼苗存活的影响, 探讨了次生针阔混交林幼苗存活影响因素及物种共存机制。结果表明: (1)适宜大树生长的局域生境同样也适宜幼苗的存活, 幼苗存活率与土壤含水量和有效氮等土壤养分显著正相关。(2)幼苗个体之间存在明显的竞争, 较多的幼苗邻体显著降低幼苗的存活率。同种大树邻体和同种幼苗邻体与幼苗存活显著负相关, 表明存在负密度制约效应。(3)随着幼苗年龄的增加, 生境异质性对幼苗存活的影响逐渐增大。该研究证实了密度制约效应和生境异质性对幼苗存活有着重要影响, 其相对重要性随着幼苗年龄级、功能群以及物种种类而变化。     

Effects of biotic neighbors and habitat heterogeneity on tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain

Aims Our study aimed to understand the effects and the relative importance of biotic neighborhood and habitat heterogeneity for tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain, north-eastern China.
Methods The generalized mixed linear model was used to examine the relative effects of biotic neighborhood and habitat heterogeneity on seedling survival over two years.
Important findings Our results showed that both biotic neighborhood and habitat heterogeneity had significant effects on the seedling survival at community level. The local environment suitable for the adult growth was also suitable for seedling survival. The soil moisture and soil available nitrogen exhibited significant positive effects on seedling survival. On the other hand, seedling density had significant negative effects on seedling survival due to the individual competition. Particularly, we found significant negative density-dependent effects on seedling survival which was caused by conspecific adult and seedling neighbors. As expected, with the increasing of seedling survival age, the habitat heterogeneity became more important on seedling survival. These results suggest that both local biotic neighborhood and habitat heterogeneity drive seedling survival in this temperate forest, and their relative importance varies with different seedling age classes and species traits.     

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.     

八大公山常绿落叶阔叶混交林影响幼苗存活的主要因子分析

幼苗是植物生活史中最脆弱的阶段,对幼苗存活影响因子的分析有助于我们更清楚的了解森林群落的天然更新机制。利用广义线性混合模型（GLMM）对八大公山常绿落叶阔叶混交林中影响幼苗存活的主要生物与非生物因子进行了研究。结果表明：（1）在群落水平上,幼苗存活与生物因子中的同种幼苗密度呈显著负相关,与非生物因子中的冠层开阔度呈显著正相关;（2）从年龄上看,4年生以下龄级的幼苗存活更容易受到同种幼苗密度的影响,与同种幼苗密度呈显著负相关;4年生及其以上的幼苗存活则主要受非生物因子影响;（3）从生活型上看,相对于常绿物种,落叶物种的幼苗存活率更容易受到同种幼苗密度的影响,也与冠层开阔度呈正相关;（4）在物种水平上,生物因子与非生物因子对不同物种幼苗存活率的影响也不相同。其中,宜昌润楠（Machilus ichangensis Rehd.et Wils.）的存活率与冠层开阔度呈正相关;薄叶山矾（Symplocos anomala Brand）幼苗的存活率与同种幼苗密度、异种大树胸高断面积、林冠开阔度、坡向均呈显著负相关,而与异种幼苗密度和海拔呈显著正相关。本研究表明影响幼苗存活的因子是多样的,而且不是随机发生的。在不同水平上影响幼苗存活的因子不同。     

The contribution of understory light availability and biotic neighborhood to seedling survival in secondary versus old-growth temperate forest

Seedling survival plays an important role in the maintenance of species diversity and forest dynamics. Although substantial gains have been made in understanding the factors driving patterns of seedling survival in forests, few studies have considered the simultaneous contribution of understory light availability and the local biotic neighborhood to seedling survival in temperate forests at different successional stages. Here, we used generalized linear mixed models to assess the relative importance of understory light availability and biotic neighborhood variables on seedling survival in secondary and old-growth temperate forest in north eastern China at two levels (community and guild). At the community level, biotic neighborhood effects on seedling survival were more important than understory light availability in both forests. In both the old-growth and secondary forests, conspecific basal area had a negative effect on seedling survival, consistent with negative conspecific density dependence. At guild levels, the relative importance of light and biotic neighborhood on seedling survival showed considerable variation among guilds in both forests. Available understory light tended to have positive effects on seedling survival for shrub and light-demanding species in the old-growth forest, but negative effects on survival of shade-tolerant seedlings in the secondary forest. For tree species and shade-tolerant species, the best fit models included neighborhood variables, but that was not the case for shrubs, light-demanding, or mid shade-tolerant species. Overall, our results demonstrate that the relative importance of understory light availability and biotic factors on seedling survival vary with species life-history strategy and forest successional stage.     

Effects of local biotic neighbors and habitat heterogeneity on seedling survival in a spruce‐fir valley forest,northeastern China

Seedlings are vulnerable to many biotic and abiotic agents, and studying seedling dynamics helps understand mechanisms of species coexistence. In this study, the relative importance of biotic neighbors and habitat heterogeneity to seedling survival was examined by generalized linear mixed models for 33 species in a spruce‐fir valley forest in northeastern China. The results showed that the relative importance of these factors varied with species and functional groups. Conspecific negative density dependence (CNDD) was important to the survival of Abies nephrolepis and Picea koraiensis seedling, whereas phylogenetic negative density dependence (PNDD) was critical to Pinus koraiensis and Betula platyphylla, as well as functional groups of tree, deciduous, and shade‐intolerant seedlings. For shrubs and Acer ukurunduense, habitat heterogeneity was significant. Despite of the significance of CNDD, PNDD, and habitat heterogeneity on seedling survival, large proportions of the total variance were not accounted for by the studied variables, suggesting the needs to examine the influences of other factors such as pests, diseases, herbivores, forest structure, species functional traits, and microclimatic conditions on seedling survival in the future.     

Density dependence across multiple life stages in a temperate old-growth forest of northeast China

Recent studies on species coexistence suggest that density dependence is an important mechanism regulating plant populations. However, there have been few studies of density dependence conducted for more than one life-history stage or that control for habitat heterogeneity, which may influence spatial patterns of survival and mask density dependence. We explored the prevalence of density dependence across multiple life stages, and the effects of controlling for habitat heterogeneity, in a temperate forest in northeast China. We used generalized linear mixed-effects models to test for density-dependent mortality of seedlings and spatial point pattern analysis to detect density dependence for sapling-to-juvenile transitions. Conspecific neighbors had a negative effect on survival of plants in both life stages. At the seedling stage, we found a negative effect of conspecific seedling neighbors on survival when analyzing all species combined. However, in species-level analyses, only 2 of 11 focal species were negatively impacted by conspecific neighbors, indicating wide variation among species in the strength of density dependence. Controlling for habitat heterogeneity did not alter our findings of density dependence at the seedling stage. For the sapling-to-juvenile transition stage, 11 of 15 focal species showed patterns of local scale (<10 m) conspecific thinning, consistent with negative density dependence. The results varied depending on whether we controlled for habitat heterogeneity, indicating that a failure to account for habitat heterogeneity can obscure patterns of density dependence. We conclude that density dependence may promote tree species coexistence by acting across multiple life-history stages in this temperate forest.     

Local neighborhood effects on long-term survival of individual trees in a neotropical forest

The survival of approximately 235 000 individual tropical trees and saplings in the 50 ha permanent plot on Barro Colorado Island (BCI), Panama was analyzed over a 13-year interval (1982–1995) as a function of four biotic neighborhood variables: (i) total stem density; (ii) conspecific density; (iii) relative plant size; and (iv) relative species richness. These neighborhood variables were measured in annular rings of width 2.5 m, extending 30 m from a given focal plant, and in one more distant annulus at 47.5–50 m. Because survival was spatially autocorrelated, a Gibbs sampler and a Monte Carlo Markov chain method were used for fitting an autologistic regression model to obtain unbiased estimates of parameter variances for hypothesis testing. After pooling all species at the community level, results showed that all four variables had significant and often strong effects on focal plant survival. Three of the four variables had negative effects on focal plant survival; relative plant size was the only variable with a positive effect (18% increase in the survival odds ratio). The variables with a negative effect on the survival odds ratio, in order of their effect strength in the nearest annulus, were: stem density (a 70% reduction in the survival odds ratio), conspecific density (50% reduction) and species richness (13% reduction). A guild-level analysis revealed considerable heterogeneity among guilds in their responses to these variables. For example, survival of gap species showed a much larger positive response to relative plant size than did survival of shade-tolerant species. Survival of shrub species was positively affected by conspecific density, but canopy tree survival was negatively affected. Conspecific density negatively affected survival of rare species much more strongly than survival of common species. The neighborhood effects of conspecific density disappear within approximately 12–15 m of the focal plant. Although locally strong, the rapid spatial decay of these effects raises unanswered questions about their quantitative contribution to the maintenance of tree diversity on landscape scales in the BCI forest.     

长白山阔叶红松林乔木幼苗组成及多度格局的影响因素

以长白山原始阔叶红松林样地为平台,以样方中胸径小于1 cm的乔木幼苗为研究对象,基于2018年两次的幼苗调查数据,利用广义线性混合模型分析影响群落乔木幼苗多度的生物邻体和生境因素。结果表明:该样地所有幼苗样方共调查到10064株乔木幼苗,累计17个物种,分属9科9属,水曲柳幼苗多度极高,在乔木树种幼苗中占优势地位;在群落水平,幼苗多度与异种大树效应、草本密度和林冠开阔度呈显著正相关,与草本盖度和土壤含水量呈显著负相关;在物种水平,水曲柳和紫椴幼苗多度的影响因素与群落水平筛选后的结果一致,红松幼苗多度与异种大树效应呈正相关,与同种大树效应及林冠开阔度呈负相关。该研究证实了生物邻体和生境因素共同影响幼苗多度格局,并且生物邻体和生境因素的相对重要性随幼苗物种种类不同而变化。     

Seedling interactions in a tropical forest in Panama

Competition is believed to be a central force limiting local diversity and controlling the structure of plant communities. However, it has been proposed that the stressed understory environment limits total understory plant density to such low levels that competitive exclusion cannot be an important factor limiting the local diversity of understory plants. To evaluate the importance of inter-seedling competition, we performed a seedling competition experiment with five shade-tolerant species in a tropical moist forest in Panama. Three-month-old seedlings were transplanted into the forest singly or with their roots intertwined with a single conspecific or heterospecific seedling in all pairwise species combinations. If competition is important, performance (survival, stem height, and number of leaves after one and six years) would be expected to be lowest with a conspecific neighbor and greatest without a neighbor. The experiment was replicated in five 0.24-m² plots at each of 20 sites in tall secondary forest. To test whether seedling performance differed among treatments we fitted linear mixed models (LMM) and generalized linear mixed models (GLMM), treating species identity and microsite (site and plot) as random effects. The five shade-tolerant study species all experienced good establishment with relatively high survival and growth rates. The neighbor treatment consistently affected seedling performance, but the effect was always very small, both in absolute terms and relative to the much stronger species and microsite effects. Seedlings with a conspecific neighbor consistently performed worse than seedlings with a heterospecific neighbor, but having no neighbor generally did not cause superior performance relative to the other treatments. We conclude that direct competitive interactions are relatively unimportant among understory plants in humid tropical forests.     

Effects of intraspecific and interspecific density on the demography of a perennial herb, Sanicula europaea

We examined the effects of intraspecific and interspecific competition on demographic processes in the perennial herb Sanicula europaea by manipulating the density of neighbouring plants. We followed the response in terms of survival, growth and reproduction and in terms of seedling recruitment. The demographic data from all phases of the life cycle enabled us to assess also the overall effects of treatments on population growth rate (λ) by transition matrix models. We also decomposed the differences in λ between control and treatments, using life table response experiments (LTRE). To study the effects of competition on recruitment in more detail and to evaluate the role of seed availability, we sowed seeds at different densities with or without vegetation removal.
Vegetative growth and flowering frequency of established individuals was not significantly affected by removal treatments, which suggest no, or a delayed response to released competition. Neighbour removal had no effect on seedling emergence but enhanced recruitment through a higher seedling survival. Conspecific and simultaneous conspecific and heterospecific removal of plants led to an increase in population growth rate (λ), whereas heterospecific removal alone led to a decrease. Emergence of seedlings and fate of vegetative established individuals contributed most to differences in λ between the control and the different treatments. Seed addition enhanced seedling emergence but, as seedling and juvenile survival were density dependent, densities of established individuals appear not to be seed limited.
In S. europaea removal treatments had different effects on established individuals and recruitment. This suggests that studies quantifying the effects of competition over the entire life cycle and performed in a natural environment are necessary to assess the importance of competition in perennial plant populations.     

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.     

Species with greater seed mass are more tolerant of conspecific neighbours: a key driver of early survival and future abundances in a tropical forest

Multiple niche‐based processes including conspecific negative density dependence (CNDD) determine plant regeneration and community structure. We ask how interspecific and intraspecific density‐dependent interactions relate to plant life histories and associated functional traits. Using hierarchical models, we analysed how such interactions affected first‐year survival of seedling recruits of 175 species in a tropical forest, and how species abundances and functional traits are related to interspecific variation in density‐dependent effects. Conspecific seedling neighbour effects prevailed over the effects of larger conspecific and all heterospecific neighbours. Tolerance of seedling CNDD enhanced recruit survival and subsequent abundance, all of which were greater among larger seeded, slow‐growing and well‐defended species. Niche differentiation along the growth–survival trade‐off and tolerance of seedling CNDD strongly correlated with regeneration success, with manifest consequences for community structure. The ability of larger seeded species to better tolerate CNDD suggests a novel mechanism for CNDD to contribute to seed‐size variation and promote species coexistence through a tolerance–fecundity trade‐off.     

Positive density dependence in seedlings of the neotropical tree species Garcinia macrophylla and Xylopia micans

Question: The Janzen‐Connell hypothesis predicts that herbivores and pathogens prevent seedlings from establishing in dense patches near adult conspecifics. Although many studies have investigated the Janzen‐Connell hypothesis, the environmental context – local or regional – in which juveniles establish is often overlooked. The objectives of this study were: (1) to evaluate Janzen‐Connell effects in contrasting environments, and (2) to incorporate microsite variation into the study of this hypothesis. Location: Pacaya‐Samiria Reserve, Peru. Methods: I assessed seedling performance of two tree species, Garcinia macrophylla and Xylopia micans, during one growing season. In an observational study, mortality and growth rates were regressed against distance to the nearest adult conspecific, conspecific seedling density, heterospecific plant density, and several abiotic variables in upland and floodplain forests. Field and shadehouse experiments were used to isolate distance‐ and density‐dependent effects. Results: Contrary to predictions, seedling survivorship increased in the presence of conspecific seedlings (Garcinia) and heterospecific understory plants (Garcinia and Xylopia) in the observational study. Survivorship in the field experiment, however, was unaffected by conspecific seedling density or adult proximity. In the shadehouse, Garcinia growth rates were highest in floodplain soils collected near adult conspecifics, but mortality was unrelated to the soil's habitat or proximity to an adult. Conclusions: The positive density dependence found in this study could have been produced by: (1) environmental factors that increase both density and survivorship, or (2) interspecific facilitation, if heterospecifics reduce herbivore or pathogen pressure on the focal species. Such interactions could help explain species coexistence in tropical forests.     

Drivers of composition and density pattern of tree seedlings in a secondary mixed conifer and broad-leaved forest,Jiaohe, Jilin,China

以吉林蛟河次生针阔混交林42 hm ²固定监测样地中209个幼苗监测站内的乔木幼苗为研究对象, 基于2016和2017年幼苗调查数据, 探究幼苗物种组成、数量动态等特征, 并运用广义线性混合效应模型分析了幼苗密度与生物邻体及其生境因素的相关性。结果表明: (1)该样地内所有幼苗监测站共调查到幼苗4 245株, 分属10科12属18种, 新生幼苗的数量在物种和调查年份间均有明显差异, 其中水曲柳(Fraxinus mandschurica)和紫椴(Tilia amurensis)幼苗表现出大量出生和死亡的动态特征。(2)群落水平上, 幼苗密度与局域同种成体胸高断面积之和、土壤全磷和有效钾等养分含量显著正相关, 适宜大树生长的生境同样适宜幼苗的生长。(3)水曲柳幼苗密度的影响因素与群落水平一致, 红松(Pinus koraiensis)存在明显的生境偏好, 在湿度较小、土壤全磷、有效氮和有效磷含量较低的生境中密度更高。相对于多年生幼苗, 同种成年邻体对当年生幼苗密度影响更加显著。该研究证实了扩散限制和生境过滤共同影响幼苗密度格局, 生物邻体和生境异质性的相对重要性随幼苗物种种类和年龄级变化。     

吉林蛟河针阔混交林乔木幼苗组成及其密度格局影响因素

以吉林蛟河次生针阔混交林42 hm ²固定监测样地中209个幼苗监测站内的乔木幼苗为研究对象, 基于2016和2017年幼苗调查数据, 探究幼苗物种组成、数量动态等特征, 并运用广义线性混合效应模型分析了幼苗密度与生物邻体及其生境因素的相关性。结果表明: (1)该样地内所有幼苗监测站共调查到幼苗4 245株, 分属10科12属18种, 新生幼苗的数量在物种和调查年份间均有明显差异, 其中水曲柳(Fraxinus mandschurica)和紫椴(Tilia amurensis)幼苗表现出大量出生和死亡的动态特征。(2)群落水平上, 幼苗密度与局域同种成体胸高断面积之和、土壤全磷和有效钾等养分含量显著正相关, 适宜大树生长的生境同样适宜幼苗的生长。(3)水曲柳幼苗密度的影响因素与群落水平一致, 红松(Pinus koraiensis)存在明显的生境偏好, 在湿度较小、土壤全磷、有效氮和有效磷含量较低的生境中密度更高。相对于多年生幼苗, 同种成年邻体对当年生幼苗密度影响更加显著。该研究证实了扩散限制和生境过滤共同影响幼苗密度格局, 生物邻体和生境异质性的相对重要性随幼苗物种种类和年龄级变化。     

Taxonomic scale-dependence of habitat niche partitioning and biotic neighbourhood on survival of tropical tree seedlings

In order to differentiate between mechanisms of species coexistence, we examined the relative importance of local biotic neighbourhood, abiotic habitat factors and species differences as factors influencing the survival of 2330 spatially mapped tropical tree seedlings of 15 species of Myristicaceae in two separate analyses in which individuals were identified first to species and then to genus. Using likelihood methods, we selected the most parsimonious candidate models as predictors of 3 year seedling survival in both sets of analyses. We found evidence for differential effects of abiotic niche and neighbourhood processes on individual survival between analyses at the genus and species levels. Niche partitioning (defined as an interaction of taxonomic identity and abiotic neighbourhood) was significant in analyses at the genus level, but did not differentiate among species in models of individual seedling survival. By contrast, conspecific and congeneric seedling and adult density were retained in the minimum adequate models of seedling survival at species and genus levels, respectively. We conclude that abiotic niche effects express differences in seedling survival among genera but not among species, and that, within genera, community and/or local variation in adult and seedling abundance drives variation in seedling survival. These data suggest that different mechanisms of coexistence among tropical tree taxa may function at different taxonomic or phylogenetic scales. This perspective helps to reconcile perceived differences of importance in the various non-mutually exclusive mechanisms of species coexistence in hyper-diverse tropical forests.     

Forest tree neighborhoods are structured more by negative conspecific density dependence than by interactions among closely related species

Interactions among neighbors influence the structure of communities of sessile organisms. Closely related species tend to share habitat and resource requirements and to interact with the same mutualists and natural enemies so that the strength of interspecific interactions tends to decrease with evolutionary divergence time. Nevertheless, the degree to which such phylogenetically related ecological interactions structure plant communities remains unclear. Using data from five large mapped forest plots combined with a DNA barcode mega‐phylogeny, we employed an individual‐based approach to assess the collective effects of focal tree size on neighborhood phylogenetic relatedness. Abundance‐weighted average divergence time for all neighbors (ADT_all) and for heterospecific neighbors only (ADT_hetero) were calculated for each individual of canopy tree species. Within local neighborhoods, we found phylogenetic composition changed with focal tree size. Specifically, significant increases in ADT_all with focal tree size were evident at all sites. In contrast, there was no significant change in ADT_hetero with tree size in four of the five sites for both sapling‐sized and all neighbors, even at the smallest neighbourhood scale (0–5 m), suggesting a limited role for phylogeny‐dependent interactions. However, there were inverse relationships between focal tree size and the proportion of heterospecific neighbors belonging to closely related species at some sites, providing evidence for negative phylogenetic density dependence. Overall, our results indicate that negative interaction with conspecifics had a much greater impact on neighborhood assemblages than interactions among closely related species and could contribute to community structure and diversity maintenance in different forest communities.