Conspecific Negative Density Dependence Research Articles

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.

Ants may buffer the Janzen-Connell effect in a tropical forest in Southwest China.

Mutualistic symbioses between ants and plants are widespread in nature. Ants can deter unwanted pests and provide protection for plants in return for food or housing rewards. Using a long-term demographic dataset in a tropical seasonal rain forest in Southwest China, we found that associations with ants positively influenced seedling survival and adult growth, and also, species with extrafloral nectaries experienced weaker conspecific negative density dependence compared with species without extrafloral nectaries. Furthermore, we found strong evidence suggesting that species in our forest experienced conspecific density dependence, which we interpreted as heavy pest pressure that may drive the development of anti-pest symbioses such as the plant-ant relationship. Our findings suggest that ants and conspecific neighbors play important but inverse roles on plant survival and growth and that ants can buffer tree neighborhood interactions in this tropical forest.

Effect of species-independent conspecific and heterospecific density dependence is contingent on seedling growth stage, season, and climate conditions

Conspecific negative density dependence (CNDD) and its extension heterospecific positive density dependence (HPDD) are the two most prominent mechanisms used to explain the maintenance of species diversity in forest communities. However, the impact of species identity, growth stage, seasonal and inter-annual variation in precipitation and temperature on conspecific and heterospecific density dependence remains poorly understood. Based on a decade of seedling (dbh < 1 cm) and tree (dbh ≥ 1 cm) census data from a 5-ha subtropical evergreen broad-leaved forest in eastern China, we used generalized linear mixed models with crossed random effects to test whether seedling survival was density dependent and varied by growth stage, species identity, season or interacted with inter-annual variation in seasonal precipitation and temperature. In addition to conspecific and heterospecific seedling and tree density, the height (or age) of the seedlings, habitat variables, seasonal precipitation and temperature and the interactions between conspecific and heterospecific seedling and tree density and seasonal precipitation and temperature were also used as predictors. The results of our long-term, spatially, and temporally explicit study showed that the strength of CNDD and enemy induced HPDD was stronger in summer and at early seedling stages, whereas interspecific facilitation induced HPDD mainly in winter for early and late seedling stages. Additionally, the effects of seasonal temperature and precipitation, and the soil PC1 axis (which correlated positively with soil nutrients and negatively with soil moisture) on seedling survival varied with season, seedling stage, and focal species identity, indicating the potential for niche partitioning among species in this community. However, neither conspecific nor heterospecific neighbor effects varied among focal seedling species, which resulted in a community compensatory trend in young seedlings. Variation in the effect of CNDD and HPDD across seedling stages, seasons, and inter-annual fluctuations of climate conditions highlights the necessity of long-term monitoring of the dynamics of CNDD in tree communities.

Disparities in tree mortality among plant functional types (PFTs) in a temperate forest: Insights into size-dependent and PFT-specific patterns

Tree mortality significantly influences forest structure and function, yet our understanding of its dynamic patterns among a range of tree sizes and among different plant functional types (PFTs) remains incomplete. This study analysed size-dependent tree mortality in a temperate forest, encompassing 46 tree species and 32,565 individuals across different PFTs (i.e., evergreen conifer vs. deciduous broadleaf species, shade-tolerant vs. shade-intolerant species). By employing all-subset regression procedures and logistic generalized linear mixed-effects models, we identified distinct mortality patterns influenced by biotic and abiotic factors. Our results showed a stable mortality pattern in evergreen conifer species, contrasted by a declining pattern in deciduous broadleaf and shade-tolerant, as well as shade-intolerant species, across size classes. The contribution to tree mortality of evergreen conifer species shifted from abiotic to biotic factors with increasing size, while the mortality of deciduous broadleaf species was mainly influenced by biotic factors, such as initial diameter at breast height (DBH) and conspecific negative density. For shade-tolerant species, the mortality of small individuals was mainly determined by initial DBH and conspecific negative density dependence, whereas the mortality of large individuals was subjected to the combined effect of biotic (competition from neighbours) and abiotic factors (i.e., convexity and pH). As for shade-intolerant species, competition from neighbours was found to be the main driver of tree mortality throughout their growth stages. Thus, these insights enhance our understanding of forest dynamics by revealing the size-dependent and PFT-specific tree mortality patterns, which may inform strategies for maintaining forest diversity and resilience in temperate forest ecosystems.

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.

Latitudinal patterns in stabilizing density dependence of forest communities

Numerous studies have shown reduced performance in plants that are surrounded by neighbours of the same species1,2, a phenomenon known as conspecific negative density dependence (CNDD)3. A long-held ecological hypothesis posits that CNDD is more pronounced in tropical than in temperate forests4,5, which increases community stabilization, species coexistence and the diversity of local tree species6,7. Previous analyses supporting such a latitudinal gradient in CNDD8,9 have suffered from methodological limitations related to the use of static data10–12. Here we present a comprehensive assessment of latitudinal CNDD patterns using dynamic mortality data to estimate species-site-specific CNDD across 23 sites. Averaged across species, we found that stabilizing CNDD was present at all except one site, but that average stabilizing CNDD was not stronger toward the tropics. However, in tropical tree communities, rare and intermediate abundant species experienced stronger stabilizing CNDD than did common species. This pattern was absent in temperate forests, which suggests that CNDD influences species abundances more strongly in tropical forests than it does in temperate ones13. We also found that interspecific variation in CNDD, which might attenuate its stabilizing effect on species diversity14,15, was high but not significantly different across latitudes. Although the consequences of these patterns for latitudinal diversity gradients are difficult to evaluate, we speculate that a more effective regulation of population abundances could translate into greater stabilization of tropical tree communities and thus contribute to the high local diversity of tropical forests.

Habitat Filtering Covers the Strength of Density Dependence and Functional Density Dependence on Seedling Survival in Cangshan Mountain, Southwest China

Conspecific negative density dependence (CNDD) is an important mechanism for species coexistence and community dynamics. Phylogenetic negative density dependence (PNDD) and functional negative density dependence (FNDD) are extensions of CNDD, and many studies have shown that they have become powerful and reliable methods for exploring the mechanisms of species coexistence. However, most studies have focused on only one or two of these mechanisms and have not considered whether and how habitat variables affect the detection of these density dependences. To investigate the relative importance of these mechanisms, we set up three 0.09 ha dynamic plots at Cangshan Mountain in southwest China, and used generalized linear mixed models to analyze how the survival of 546 woody plant seedlings was affected by neighborhood density and habitat variables. Our results showed that heterospecific seedling density dependence and functional trait density dependence played key roles in seedling survival. Habitat factors, phylogenetic densities, and adult neighbors had no significant effect on seedling survival in the three plots. However, habitat filtering covered the detection of density dependence and functional trait density dependence. Our study demonstrates that failure to control for habitat variables may obscure the importance of density dependence and functional trait density dependence on seedling survival.

Mycorrhizal feedbacks influence global forest structure and diversity

One mechanism proposed to explain high species diversity in tropical systems is strong negative conspecific density dependence (CDD), which reduces recruitment of juveniles in proximity to conspecific adult plants. Although evidence shows that plant-specific soil pathogens can drive negative CDD, trees also form key mutualisms with mycorrhizal fungi, which may counteract these effects. Across 43 large-scale forest plots worldwide, we tested whether ectomycorrhizal tree species exhibit weaker negative CDD than arbuscular mycorrhizal tree species. We further tested for conmycorrhizal density dependence (CMDD) to test for benefit from shared mutualists. We found that the strength of CDD varies systematically with mycorrhizal type, with ectomycorrhizal tree species exhibiting higher sapling densities with increasing adult densities than arbuscular mycorrhizal tree species. Moreover, we found evidence of positive CMDD for tree species of both mycorrhizal types. Collectively, these findings indicate that mycorrhizal interactions likely play a foundational role in global forest diversity patterns and structure.

Modification of Density Dependence and Habitat Filtering on Seedling Survival of Different Mycorrhizal-Type Tree Species in Temperate Forests

Conspecific negative density dependence (CNDD) and habitat filtering are critical to seedling survival. However, the relative importance of the two processes in affecting survival of seedlings with different types of mycorrhizae remains unclear. In this study, the effects of CNDD and habitat filtering on the survival of tree seedlings with different mycorrhizal types were investigated at different successional stages of a temperate forest in the Changbai Mountain Natural Reserve, Northeast China. Conspecific negative density dependence and habitat filtering significantly affected seedling survival. In the early successional stage, the interactions between conspecific neighbor tree density and light availability and soil properties significantly negatively affected survival of all species and arbuscular mycorrhizal (AM) seedlings in the community, but not that of ectomycorrhizal (EcM) seedlings, and the CNDD effect was stronger on AM seedlings than on EcM seedlings. In the mid-successional stage, CNDD effects were stronger on EcM seedlings. Therefore, different types of mycorrhizal seedlings responded differently to CNDD and habitat filtering mechanisms during community succession, and thus, tree mycorrhizal association could determine the effects of CNDD and habitat filtering on seedling survival in temperate forests.

Pervasive within-species spatial repulsion among adult tropical trees.

For species to coexist, performance must decline as the density of conspecific individuals increases. Although evidence for such conspecific negative density dependence (CNDD) exists in forests, the within-species spatial repulsion it should produce has rarely been demonstrated in adults. In this study, we show that in comparison to a null model of stochastic birth, death, and limited dispersal, the adults of dozens of tropical forest tree species show strong spatial repulsion, some to surprising distances of approximately 100 meters. We used simulations to show that such strong repulsion can only occur if CNDD considerably exceeds heterospecific negative density dependence-an even stronger condition required for coexistence-and that large-scale repulsion can indeed result from small-scale CNDD. These results demonstrate substantial niche differences between species that may stabilize species diversity.

Age, size and neighbors influence the survival and growth of understory trees in a naturally reproducing population of American chestnut, Castanea dentata

The seedling stage is a critical point in the life history of plants. The survival and growth of tree seedlings can be affected by both the abiotic and biotic environment. Seedlings of both tropical and temperate forests have been shown to experience conspecific negative density dependence (CNDD) and positive density dependence (facilitation). American chestnut was once a common species throughout eastern North America that was decimated by an introduced pathogen in the early 1900 s. Here we present an analysis of chestnut seedling ecology in a unique, sexually reproducing population of American chestnut in central Maine, USA, to understand how both conspecific and heterospecific canopy trees along with size and age influence the survival and growth of understory chestnut trees. Using both a cohort analysis and a static life table approach, we found that seedling survival was lower in the first year of life, and those trees that survive their first and second years had consistently high survival. Our data did not support facilitation as none of the neighborhood indices were positive. We found evidence for CNDD for American chestnuts in the first year of life, but we found no evidence for continued effects beyond the first year. Heterospecifics decreased growth and survival of understory chestnuts beyond the first year of life. Enhanced understanding of American chestnut ecology, particularly at the critical seedling stage, can guide management strategies that support reintroduction efforts.

Trees species’ dispersal mode and habitat heterogeneity shape negative density dependence in a temperate forest

Conspecific negative density dependence (CNDD) is a potentially important mechanism in maintaining species diversity. While previous evidence showed habitat heterogeneity and species’ dispersal modes affect the strength of CNDD at early life stages of trees (e.g., seedlings), it remains unclear how they affect the strength of CNDD at later life stages. We examined the degree of spatial aggregation between saplings and trees for species dispersed by wind and gravity in four topographic habitats within a 25-ha temperate forest dynamic plot in the Qinling Mountains of central China. We used the replicated spatial point pattern (RSPP) analysis and bivariate pair-correlation function (PCF) to detect the spatial distribution of saplings around trees at two scales, 15 and 50 ​m, respectively. Although the signal was not apparent across the whole study region (or 25-ha), it is distinct on isolated areas with specific characteristics, suggesting that these characteristics could be important factors in CNDD. Further, we found that the gravity-dispersed tree species experienced CNDD across habitats, while for wind-dispersed species CNDD was found in gully, terrace and low-ridge habitats. Our study suggests that neglecting the habitat heterogeneity and dispersal mode can distort the signal of CNDD and community assembly in temperate forests.

Effects of Disturbance and Conspecific Negative Density Dependence on Forest Composition and Diversity: A Simulation-Based Approach

Effects of Disturbance and Conspecific Negative Density Dependence on Forest Composition and Diversity: A Simulation-Based Approach

Coexistence is stabilized by conspecific negative density dependence via fungal pathogens more than oomycete pathogens.

Plant pathogens are often hypothesized to promote species coexistence by generating conspecific negative density dependence (CNDD). However, the relative importance of fungal versus oomycete pathogens in maintaining plant species coexistence and community composition remains unresolved, despite their recognized effects on plant performance. Here, we use fungicide application to investigate how fungal versus oomycete pathogens affect plant species coexistence in an alpine meadow. We found that the severity of foliar fungal disease was density-dependent at both intra- and interspecific levels. Fungal pathogen-exclusion treatment successfully decreased the severity of foliar fungal diseases, with no detectable effects on root colonization by arbuscular mycorrhizal fungi or on soil chemical properties. Fungal pathogens were important factors shaping CNDD across 25 coexisting plant species. Exclusion of fungal pathogens significantly reduced plant species richness and Shannon's evenness. Treatments that excluded fungal pathogens also led to significant shifts in plant community composition toward more Poaceae and Cyperaceae. These results indicate that fungal pathogens, especially those affecting aboveground plant parts, may play a larger role in maintaining species coexistence and shaping community composition than has been previously recognized.

Differential impacts of adult trees on offspring and non-offspring recruits in a subtropical forest.

An important mechanism promoting species coexistence is conspecific negative density dependence (CNDD), which inhibits conspecific neighbors by accumulating host-specific enemies near adult trees. Natural enemies may be genotype-specific and regulate offspring dynamics more strongly than non-offspring, which is often neglected due to the difficulty in ascertaining genetic relatedness. Here, we investigated whether offspring and non-offspring of a dominant species, Castanopsis eyrei, suffered from different strength of CNDD based on parentage assignment in a subtropical forest. We found decreased recruitment efficiency (proxy of survival probability) of offspring compared with non-offspring near adult trees during the seedling-sapling transition, suggesting genotype-dependent interactions drive tree demographic dynamics. Furthermore, the genetic similarity between individuals of same cohort decreased in late life history stages, indicating genetic-relatedness-dependent tree mortality throughout ontogeny. Our results demonstrate that within-species genetic relatedness significantly affects the strength of CNDD, implying genotype-specific natural enemies may contribute to population dynamics in natural forests.

Differences in Density Dependence among Tree Mycorrhizal Types Affect Tree Species Diversity and Relative Growth Rates.

Conspecific negative density dependence (CNDD) may vary by tree mycorrhizal type. However, whether arbuscular mycorrhizal (AM)-associated tree species suffer from stronger CNDD than ectomycorrhizal (EcM) and ericoid mycorrhizal (ErM)-associated tree species at different tree life stages, and whether EcM tree species can promote AM and ErM saplings and adults growth, remain to be studied. Based on the subtropical evergreen broad-leaved forest data in eastern China, the generalized linear mixed-effects model was used to analyze the effects of the conspecific density and heterospecific density grouped by symbiont mycorrhizal type on different tree life stages of different tree mycorrhizal types. The results showed that compared to other tree mycorrhizal types at the same growth stage, EcM saplings and AM adults experienced stronger CNDD. Heterospecific EcM density had a stronger positive effect on AM and ErM individuals. Species diversity and average relative growth rate (RGR) first increased and then decreased with increasing basal area (BA) ratios of EcM to AM tree species. These results suggested that the stronger CNDD of EcM saplings and AM adults favored local species diversity over other tree mycorrhizal types. The EcM tree species better facilitated the growth of AM and ErM tree species in the neighborhood, increasing the forest carbon sink rate. Interestingly, species diversity and average RGR decreased when EcM or AM tree species predominated. Therefore, our study highlights that manipulating the BA ratio of EcM to AM tree species will play a nonnegligible role in maintaining biodiversity and increasing forest carbon sink rates.

Disentangling the effects of species interactions and environmental factors on the spatial pattern and coexistence of two congeneric Pinus species in a transitional climatic zone.

Congeneric species are critical for understanding the underlying ecological mechanisms of biodiversity maintenance. Ecological mechanisms such as conspecific negative density dependence, species differences in life‐history stages related to habitat preference, and limiting similarity are known to influence plant fitness, thereby influencing species coexistence and biodiversity. However, our understanding of these phenomena as they apply to coexistence among coniferous species is limited. We studied two congeneric Pinus species, Pinus armandii (PA) and Pinus tabulaeformis (PT), both of which are common pioneer species typically succeeded by oaks (Quercus), in a 25‐ha warm temperate deciduous broad‐leaved forest. Here, we addressed the following questions: (1) How do population structures and distributions patterns of these two Pinus species vary with respect to different life‐history stages? (2) Does intra‐ and interspecific competition vary with respect to three life‐history stages? And (3) What are the relative contributions of topographic and soil variables to the spatial distributions of the species across the three life‐history stages? In addressing these questions, we utilized the pair‐correlation function g(r), redundancy analysis (RDA), variance partitioning (VP), and hierarchical partitioning (HP) to identify habitat preferences and conspecific negative density dependence at different life‐history stages from small to large trees. The results revealed that in both Pinus species, individuals in different life‐history stages were subject to significant habitat heterogeneity, with a tendency for small trees to be distributed at higher latitudes that may be represents climate‐change‐driven migration in both species. In addition, the effects of conspecific negative density dependence on PT were stronger than those on PA due to limited dispersal in PT. Furthermore, we found that interspecific competition was weak due to the species differences in resource utilization and preference for key habitats. Our study shows that congeneric Pinus species avoids competition by exploiting distinct habitats and provides insight into forest community structure.

Experimental and observational evidence of negative conspecific density dependence in temperate ectomycorrhizal trees.

Conspecific negative density dependence (CNDD) promotes tree species diversity by reducing recruitment near conspecific adults due to biotic feedbacks from herbivores, pathogens, or competitors. While this process is well-described in tropical forests, tests of temperate tree species range from strong positive to strong negative density dependence. To explain this, several studies have suggested that tree species traits may help predict the strength and direction of density dependence: for example, ectomycorrhizal-associated tree species typically exhibit either positive or weaker negative conspecific density dependence. More generally, the strength of density dependence may be predictably related to other species-specific ecological attributes such as shade tolerance, or the relative local abundance of a species. To test the strength of density dependence and whether it affects seedling community diversity in a temperate forest, we tracked the survival of seedlings of three ectomycorrhizal-associated species experimentally planted beneath conspecific and heterospecific adults on the Prospect Hill tract of the Harvard Forest, in Massachusetts, USA. Experimental seedling survival was always lower under conspecific adults, which increased seedling community diversity in one of six treatments. We compared these results to evidence of CNDD from observed sapling survival patterns of 28 species over approximately 8 years in an adjacent 35-ha forest plot. We tested whether species-specific estimates of CNDD were associated with mycorrhizal association, shade tolerance, and local abundance. We found evidence of significant, negative conspecific density dependence (CNDD) in 23 of 28 species, and positive conspecific density dependence in two species. Contrary to our expectations, ectomycorrhizal-associated species generally exhibited stronger (e.g., more negative) CNDD than arbuscular mycorrhizal-associated species. CNDD was also stronger in more shade-tolerant species but was not associated with local abundance. Conspecific adult trees often have a negative influence on seedling survival in temperate forests, particularly for tree species with certain traits. Here we found strong experimental and observational evidence that ectomycorrhizal-associating species consistently exhibit CNDD. Moreover, similarities in the relative strength of density dependence from experiments and observations of sapling mortality suggest a mechanistic link between negative effects of conspecific adults on seedling and sapling survival and local tree species distributions.

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.

Interannual climate variability has predominant effects on seedling survival in a temperate forest.

Mechanisms such as conspecific negative density dependence (CNDD) and niche partitioning have been proposed to explain species coexistence and community diversity. However, as a potentially important axis of niche partitioning, the role of interannual climate variability in driving local community dynamics remains largely unknown. Here we used a 15-year monitoring data set of more than 53,000 seedlings in a temperate forest to examine (1) what are the relative effects of interannual climate variability, biotic interactions, and habitat conditions on seedling survival; (2) how the effects of biotic interactions change with interannual climate variability, and habitat conditions; and (3) whether the impacts of interannual climate variability, biotic interactions, and habitat conditions differ with plant traits. Interannual climate variability accounted for the most variation in seedling survival at the community level, followed by biotic interactions, and habitat conditions. Increased snowpack and decreased minimum temperature during the non-growing season had positive effects on seedling survival. Effects of conspecific neighbor density were weakened in higher snowpack, effective accumulated temperature, elevation, and soil-resource gradient, but were intensified with increased ultraviolet radiation, maximum precipitation, minimum temperature, and soil moisture. In addition, the relative importance of interannual climate variability versus biotic interactions differed depending on species-trait groups. Specifically, biotic interactions for gravity-dispersed species had a larger effect size in affecting seedling survival than other trait groups. Also, gravity-dispersed species experienced a stronger CNDD than wind-dispersed, probably because wind-dispersed seedlings rarely had adult conspecifics nearby. We found that interannual climate variability was most strongly associated with seedling survival, but the magnitude of climatic effects varied among species-trait groups. Interannual climate variability may act as an inhibitor or accelerator to density-dependent interactions and should be accounted for in future studies, as both a potential direct and indirect factor in understanding the diversity of forest communities.