Explain Species Coexistence Research Articles

Characterization of Invasiveness, Thermotolerance and Light Requirement of Nine Invasive Species in China

Understanding responsible functional traits for promoting plant invasiveness could be important to aid in the development of adequate management strategies for invasive species. Seed traits play an important role in the plant life cycle by affecting dispersal ability, formation of the soil seed bank, type and level of dormancy, germination, survival and/or competitive ability. We assessed seed traits and germination strategies of nine invasive species under five temperature regimes and light/dark treatments. Our results showed a considerable level of interspecific variation in germination percentage among the tested species. Both cooler (5/10 °C) and warmer (35/40 °C) temperatures tended to inhibit germination. All study species were considered small-seeded, and seed size did not affect germination in the light. Yet, a slightly negative correlation was found between germination in the dark and seed dimensions. We classified the species into three categories according to their germination strategies: (i) risk-avoiders, mostly displaying dormant seeds with low G%; (ii) risk-takers, reaching a high G% in a broad range of temperatures; (iii) intermediate species, showing moderate G% values, which could be enhanced in specific temperature regimes. Variability in germination requirements could be important to explain species coexistence and invasion ability of plants to colonize different ecosystems.

A trade-off between space exploration and mobilization of organic phosphorus through associated microbiomes enables niche differentiation of arbuscular mycorrhizal fungi on the same root.

Ecology seeks to explain species coexistence, but experimental tests of mechanisms for coexistence are difficult to conduct. We synthesized an arbuscular mycorrhizal (AM) fungal community with three fungal species that differed in their capacity of foraging for orthophosphate (P) due to differences in soil exploration. We tested whether AM fungal species-specific hyphosphere bacterial assemblages recruited by hyphal exudates enabled differentiation among the fungi in the capacity of mobilizing soil organic P (Po). We found that the less efficient space explorer, Gigaspora margarita, obtained less 13C from the plant, whereas it had higher efficiencies in Po mobilization and alkaline phosphatase (AlPase) production per unit C than the two efficient space explorers, Rhizophagusintraradices and Funneliformis mosseae. Each AM fungus was associated with a distinct alp gene harboring bacterial assemblage, and the alp gene abundance and Po preference of the microbiome associated with the less efficient space explorer were higher than those of the two other species. We conclude that the traits of AM fungal associated bacterial consortia cause niche differentiation. The trade-off between foraging ability and the ability to recruit effective Po mobilizing microbiomes is a mechanism that allows co-existence of AM fungal species in a single plant root and surrounding soil habitat.

Using leaf traits to explain species co-existence and its consequences for primary productivity across a forest-steppe ecotone

Trait-based approaches have been widely applied to uncover the mechanisms determining community assembly and biodiversity-ecosystem functioning relationships. However, they have rarely been used in forest-steppe ecotones. These ecosystems are extremely sensitive to disturbances due to their relatively complex ecosystem structures, functionings and processes. In this study, we selected seven sites along a transect from closed canopy forests (CF) to forest-steppe ecotones (FSE) and meadow steppes (MS) in northeast China. Six leaf functional traits (i.e. leaf nitrogen and phosphorus contents, leaf length and thickness, single leaf area and leaf mass per unit area, LMA) as well as the community composition and aboveground biomass at each site were measured. Both functional trait diversity indices (richness, evenness and divergence) and community-weighted mean trait values (CWMs) were calculated to quantify community trait distributions. We found that dominant species in the FSE communities showed acquisitive strategies with highest leaf nitrogen (Mean ± SE: 19.6 ± 0.5 mg g−1) and single leaf area (19.2 ± 1.3 cm2), but the lowest LMA (59.6 ± 1.3 g cm−2) values compared to adjacent CF and MS communities. The ecotone communities also exhibited the largest functional trait richness (TOP), evenness (TED) and divergence (FDis) values (0.46, 0.92 and 0.67, respectively). Overall, niche differentiation emerges as the main mechanism influencing the coexistence of plant species in ecotone ecosystems. In addition, CWMs of leaf traits were the most important predictors for estimating variations in aboveground productivity across the transect, suggesting a major influence of dominant species. Our findings suggest that vegetation management practices in forest-steppe ecotones should increasingly focus on community functional trait diversity, and support the establishment and regeneration of plant species with rapid resource acquisition strategies.

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.

Seed dispersal by wind decreases when plants are water-stressed, potentially counteracting species coexistence and niche evolution.

Hydrology is a major environmental factor determining plant fitness, and hydrological niche segregation (HNS) has been widely used to explain species coexistence. Nevertheless, the distribution of plant species along hydrological gradients does not only depend on their hydrological niches but also depend on their seed dispersal, with dispersal either weakening or reinforcing the effects of HNS on coexistence. However, it is poorly understood how seed dispersal responds to hydrological conditions. To close this gap, we conducted a common‐garden experiment exposing five wind‐dispersed plant species (Bellis perennis, Chenopodium album, Crepis sancta, Hypochaeris glabra, and Hypochaeris radicata) to different hydrological conditions. We quantified the effects of hydrological conditions on seed production and dispersal traits, and simulated seed dispersal distances with a mechanistic dispersal model. We found species‐specific responses of seed production, seed dispersal traits, and predicted dispersal distances to hydrological conditions. Despite these species‐specific responses, there was a general positive relationship between seed production and dispersal distance: Plants growing in favorable hydrological conditions not only produce more seeds but also disperse them over longer distances. This arises mostly because plants growing in favorable environments grow taller and thus disperse their seeds over longer distances. We postulate that the positive relationship between seed production and dispersal may reduce the concentration of each species to the environments favorable for it, thus counteracting species coexistence. Moreover, the resulting asymmetrical gene flow from favorable to stressful habitats may slow down the microevolution of hydrological niches, causing evolutionary niche conservatism. Accounting for context‐dependent seed dispersal should thus improve ecological and evolutionary models for the spatial dynamics of plant populations and communities.

Acoustic segregation of five sympatric and syntopic species of genus Pristimantis (Anura: Strabomantidae) from Western Colombia

ABSTRACT The Andean Cordilleras of Colombia, especially the Cordillera Occidental, are among the areas with the highest diversity of Pristimantis frogs in the world. Within the Cordillera Occidental, the Serranía de los Paraguas is famous for its high diversity of sympatric species of Pristimantis frogs. In this study, we investigated acoustic frequency segregation in five sympatric and syntopic species of the genus Pristimantis inhabiting this Serranía. It is important to point out that three of the five species’ calls are described for the first time: Pristimantis brevifrons, P. ptochus, and P. silverstonei. Our results showed that acoustic frequency segregation occurs among these five Pristimantis species. Another notable result was the inverse relationship between dominant frequency and male body size, which was corroborated in all species except in P. alius. Finally, we suggested that for improving our knowledge on species coexistence in Pristimantis, future research should also focus on species history and character evolution, given that the evolutionary history is also fundamental for explaining species coexistence.

Limited evidence of coupling between above and belowground functional traits in tropical dry forest seedlings

Introduction: Water availability is one of the main factors determining the distribution of woody species in the tropics. Although the functional mechanisms that determine the species tolerance to water deficit have been extensively studied in adult individuals, the responses of early ontogenetic stages have been less explored. Objective: To identify functional strategies and trait correlations between different seedlings' dimensions (leaf, stem, and root). We expect limited coordination between above and below-ground functional traits due to a single conservation-acquisition trade-off cannot capture the variability of functions and environmental pressures to which the root system is subjected. Methods: We measured 12 functional traits belonging to 38 seedling species in a tropical dry forest in Colombia. We explored the relationships between pairs of traits using Pearson correlations, and to obtain an integrated view of the functional traits, a principal component analysis (PCA) was performed. Results: The results showed limited evidence of linkage between above- and below-ground traits, but we did find significant correlations between traits for the continuum of conservative and acquisitive strategies. Root traits related to water and nutrient take capacity formed an orthogonal axis to the acquisitive-conservative continuum. Conclusions: Our results showed that dry forest seedlings have different functional strategies to cope with water deficit. The incorporation of root traits helps to explain new functional strategies not reported for leaf and stem traits. This study contributes to understanding the mechanisms that explain species coexistence and is particularly relevant for predicting future forest trajectories.

Plant Community Assembly Mechanisms of a Subalpine Meadow Community Along Different Successional Time

The process of community assembly is important to explain species coexistence and the maintenance of species diversity. Succession provides a good opportunity to study the role of the deterministic and stochastic processes during the community species composition. We investigated species composition and environmental factors and calculated the change in species diversity, phylogenetic diversity, net relatedness index, and nearest taxon index along a successional chronosequence, with a series of 0.5 × 0.5 m2 plots (n = 20) from five successional subalpine meadow plant communities (ages 1, 3, 5, 15, and 30) in the Qinghai-Tibetan Plateau. In the plots, we found a significant change in species composition and abiotic environmental factors. Our redundancy analysis showed that soil nitrate nitrogen and soil organic carbon were the key environmental factors that affect the community composition in succession gradient. The community showed an overdispersion phylogenetic structure along with the early and medium succession. The ends of the succession stage showed a random or convergent phylogenetic structure. The results showed that the construction process of plant communities in the early and middle stages of succession was mainly affected by habitat filtering. In the late stage of succession, the process of community construction is more complex, which could be affected by the limiting similarity and neutral driving process. Our findings are important to be used in managing ecological restoration and protection.

Functional seed traits and germination patterns predict species coexistence in Northeast Mediterranean foredune communities.

The structure of plant communities, which is based on species abundance ratios, is closely linked to ecosystem functionality. Seed germination niche plays a major role in shaping plant communities, although it has often been neglected when explaining species coexistence. The aim of this work is to link the seed germination niche to community ecology, investigating how functional seed traits contribute to species coexistence. Species selection was based on a database of 504 vegetation surveys from the Veneto coast (Italy). Through cluster analysis we identified the foredune community and selected all of its 19 plant species. By using the 'Phi coefficient' and frequency values, species were pooled in different categories (foundation species, accidental species of the semi-fixed dune and aliens), then the 19 species were grouped according to their germination responses to temperature and photoperiod through cluster analyses. For each germination cluster, we investigated germination trends against temperature and photoperiod by using generalized linear mixed models. We identified four germination strategies: (1) high germination under all tested conditions ('high-germinating'); (2) high germination at warm temperatures in the dark ('dark warm-cued'); (3) high germination at warm temperatures in the light ('light warm-cued'); and (4) low germination, regardless of conditions ('low-germinating'). Foredune foundation species showed a narrow germination niche, being 'low-germinating' or 'dark warm-cued'. Annual species of semi-fixed dunes were 'high-germinating', while alien species were the only members of the 'light warm-cued' cluster. Our research suggests that different categories of species have dissimilar seed germination niches, which contributes to explaining their coexistence. Climatic events, such as rising temperature, could alter germination patterns, favouring seed regeneration of certain categories (i.e. alien and semi-fixed dune species) at the expense of others (i.e. foundation species, pivotal to ecosystem functioning), and hence potentially altering the plant community structure.

Hydrologic niches explain species coexistence and abundance in a shrub–steppe system

Abstract Differences in vertical root distributions are often assumed to create resource uptake trade‐offs that determine plant growth and coexistence. Yet, most plant roots are in shallow soils, and data linking root distributions with resource uptake and plant abundances remain elusive. Here we used a tracer experiment to describe the vertical distribution of absorptive roots of dominant species in a shrub–steppe ecosystem. To describe how these different rooting distributions affected water uptake in wet and dry soils across a growing season, we used a soil water movement model. Root traits were then correlated with plant landscape abundances. Deeper root distributions extracted more soil water, had larger unique hydrological niches and were more abundant on the landscape. Though most (>50%) root biomass and tracer uptake occurred in shallow soils (0–32 cm), the depth of 50% of tracer uptake varied from 11 to 32 cm across species and species with deeper rooting distributions were more abundant on the landscape (R2 = .95). The water flow model revealed that deeper rooting distributions should extract more soil water (i.e. a range of 60–113 mm of soil water) because shallow roots were often in dry soils. These potential water uptake values were tightly correlated with species’ abundances on the landscape (R2 = .90). Finally, each species’ rooting distribution demonstrated a depth and time at which it could extract more soil water than any other rooting distribution, and the size of these unique hydrological niches indices was also well correlated with species’ abundances (R2 = .89). Synthesis. Our results demonstrate not only a correlation between root distributions and species abundance, but also the mechanism through which differences in rooting distributions can determine resource uptake and niche partitioning, even when most roots are found in shallow soils.

Arboreal twig-nesting ants form dominance hierarchies over nesting resources.

Interspecific dominance hierarchies have been widely reported across animal systems. High-ranking species are expected to monopolize more resources than low-ranking species via resource monopolization. In some ant species, dominance hierarchies have been used to explain species coexistence and community structure. However, it remains unclear whether or in what contexts dominance hierarchies occur in tropical ant communities. This study seeks to examine whether arboreal twig-nesting ants competing for nesting resources in a Mexican coffee agricultural ecosystem are arranged in a linear dominance hierarchy. We described the dominance relationships among 10 species of ants and measured the uncertainty and steepness of the inferred dominance hierarchy. We also assessed the orderliness of the hierarchy by considering species interactions at the network level. Based on the randomized Elo-rating method, we found that the twig-nesting ant species Myrmelachista mexicana ranked highest in the ranking, while Pseudomyrmex ejectus was ranked as the lowest in the hierarchy. Our results show that the hierarchy was intermediate in its steepness, suggesting that the probability of higher ranked species winning contests against lower ranked species was fairly high. Motif analysis and significant excess of triads further revealed that the species networks were largely transitive. This study highlights that some tropical arboreal ant communities organize into dominance hierarchies.

Spatial patterns and associations between species belonging to four genera of the Lauraceae family.

Spatial distribution pattern of biological related species present unique opportunities and challenges to explain species coexistence. In this study, we explored the spatial distributions and associations among congeneric species at both the species and genus levels to explain their coexistence through examining the similarities and differences at these two levels. We first used DNA and cluster analysis to confirmed the relative relationship of eight species within a 20 ha subtropical forest in southern China. We compared Diameter at breast height (DBH) classes, aggregation intensities and spatial patterns, associations, and distributions of four closely related species pairs to reveal similarities and differences at the species and genus levels. These comparisons provided insight into the mechanisms of coexistence of these congeners. O-ring statistics were used to measure spatial patterns of species. Ω 0–10, the mean conspecific density within 10 m of a tree, was used as a measure of the intensity of aggregation of a species, and g-function was used to analyze spatial associations. Our results suggested that spatial aggregations were common, but the differences between spatial patterns were reduced at the genus level. Aggregation intensity clearly reduced at the genus level. Negative association frequencies decreased at the genus level, such that independent association was commonplace among all four genera. Relationships between more closely related species appeared to be more competitive at both the species and genus levels. The importance of competition on interactions is most likely influenced by similarity in lifestyle, and the habitat diversity within the species’ distribution areas. Relatives with different lifestyles likely produce different distribution patterns through different interaction process. In order to fully understand the mechanisms generating spatial distributions of coexisting siblings, further research is required to determine the spatial patterns and associations at other classification levels.

Rare species occupy uncommon niches.

The fact that temperate grasslands often contain upwards of 30 vascular plant species per m2 yet these species seem to have relatively similar life histories and resource requirements has made explaining species coexistence in these communities a major focus of research. While the reduction of competition by disturbance has been a popular explanation for species coexistence, in tallgrass prairies any level of disturbance either has no effect, or decreases diversity, since it favors the dominant plants. Although there has long been speculation that grassland species could coexist by niche partitioning the concept received renewed interest when it was shown that soil hydrology could explain species coexistence. One aspect of community structure that has not been explained by niche partitioning is the rareness and commonness of species within communities. There are three classes of explanations for rareness: narrow habitat requirements, low competitive ability combined with frequency dependent fitness and, dispersal ability. However, evidence for these explanations tend to be anecdotal, focusing on particular species. Here I show that in tallgrass prairies common and rare species consistently occupy different parts of niche space, with rare species being restricted by the cover of common species and occupying the rare available niches.

Spatial patterns and associations of four species in an old-growth temperate forest

The spatial pattern of a tree species is an important characteristic of plant communities, providing critical information to explain species coexistence. The spatial distribution and association of four different successional species were analyzed among different life-history stages in an old-temperate forest. Significant aggregation patterns were found, and the degree of aggregation decreased with the scales and life-history stages. Significant interspecific spatial associations were detected. In comparing the relationships among the different life-history stages, positive associations were found at small scales in all of the juvenile species pairs. In the adult stage, negative associations were detected in coniferous vs. deciduous species pairs, while the deciduous species pairs, which have identical resource requirements, showed a positive association in this study. The coniferous species pairs showed a positive association at small scales. We infer that seed dispersal, competitive ability, or the requirement for specific topographic and light environments may contribute to the coexistence of these species.

Dynamics and coexistence in a system with intraguild mutualism

It is a tenet of ecological theory that two competing consumers cannot stably coexist on a single limiting resource in a homogeneous environment. Many mechanisms and processes have since been evoked and studied, empirically and theoretically, to explain species coexistence and the observed biological diversity. Facilitative interactions clearly have the potential to enhance coexistence. Yet, even though mutual facilitation between species of the same guild is widely documented empirically, the subject has received very little theoretical attention. Here, we study one form of intraguild mutualism in the simplest possibly community module of one resource and two consumers. We incorporate mutualism as enhanced consumption in the presence of the other consumers. We find that intraguild mutualism can (a) significantly enhance coexistence of consumers, (b) induce cyclic dynamics, and (c) give rise to a bi-stability (a ‘joint’ Allee effect) and potentially catastrophic collapse of both consumer species.

Dispersal-mediated coexistence under recruitment limitation and displacement competition

Understanding biodiversity maintenance when species compete for shared limiting resources remains an outstanding ecological problem. For a half century, the competition–colonization trade-off has been invoked to explain species coexistence. More recently, asymmetric dispersal has been shown to alter the conditions for species coexistence under pure scramble competition for space, even in the absence of spatial heterogeneity. Therefore, in this study, we investigated why and how asymmetric dispersal may alter the effect of competition–colonization trade-off on species coexistence. Our main findings are that: (1) self-recruitment of a superior competitor facilitates persistence of an inferior competitor; (2) the degree to which a community is recruitment-limited influences the threshold dispersal asymmetry required for persistence of a species with a birth or survival disadvantage; (3) an inferior competitor with a high recruitment strategy is better able to benefit from spatial asymmetries in dispersal than one with a high survival strategy, and (4) an inferior competitor may persist, and even exclude the superior competitor, when dispersal is highly asymmetric (i.e., the competition–colonization trade-off can be violated). These results underscore the importance of dispersal patterns promoting species coexistence and their synergies with species’ colonization and competition abilities, and highlight the need to assess dispersal-mediated coexistence within a framework that incorporates the interactive effects of dispersal patterns, demographic and competitive strategies.

Competition–colonization dynamics in experimental bacterial metacommunities

One of the simplest hypotheses used to explain species coexistence is the competition-colonization trade-off, that is, species can stably coexist in a landscape if they show a trade-off between competitive and colonization abilities. Despite extensive theory, the dynamics predicted to result from competition-colonization trade-offs are largely untested. Landscape change, such as habitat destruction, is thought to greatly influence coexistence under competition-colonization dynamics, although there is no formal test of this prediction. Here we present the first illustration of competition-colonization dynamics that fully transposes theory into a controlled experimental metacommunity of two Pseudomonas bacterial strains. The competition-colonization dynamics were achieved by directly manipulating trade-off strength and colonization rates to generate the full range of coexistence conditions and responses to habitat destruction. Our study successfully generates competition-colonization dynamics matching theoretical predictions, and our results further reveal a negative relationship between diversity and productivity when scaling up to entire metacommunities.

Emergent neutrality drives phytoplankton species coexistence.

The mechanisms that drive species coexistence and community dynamics have long puzzled ecologists. Here, we explain species coexistence, size structure and diversity patterns in a phytoplankton community using a combination of four fundamental factors: organism traits, size-based constraints, hydrology and species competition. Using a 'microscopic' Lotka-Volterra competition (MLVC) model (i.e. with explicit recipes to compute its parameters), we provide a mechanistic explanation of species coexistence along a niche axis (i.e. organismic volume). We based our model on empirically measured quantities, minimal ecological assumptions and stochastic processes. In nature, we found aggregated patterns of species biovolume (i.e. clumps) along the volume axis and a peak in species richness. Both patterns were reproduced by the MLVC model. Observed clumps corresponded to niche zones (volumes) where species fitness was highest, or where fitness was equal among competing species. The latter implies the action of equalizing processes, which would suggest emergent neutrality as a plausible mechanism to explain community patterns.

Assessing habitat differentiation between coexisting species: The role of spatial scale

Although the importance of spatial scale for understanding habitat selection patterns and processes has long been recognized, little is known about its impact on the identification of habitat differentiation between sympatric species, despite its likely utility in assessing niche partitioning and thus explaining species coexistence. Here we used radio-telemetry data to examine seasonal habitat selection and differentiation by Cabrera ( n = 28) and water voles ( n = 29) within habitat patches in a highly fragmented landscape, across spatial extents and resolutions. Habitat selection was found for both species at the home-range and core-area scales, tending to be strongest for water and Cabrera voles at coarse and fine spatial resolutions, respectively. Water voles showed higher preference for humid sedge/rush and reed habitats across seasons and spatial scales. Cabrera voles consistently selected tall grass and shrub habitats during the wet season, whereas dry season preference was higher for sedge/rush and tall grass at fine and coarse spatial resolutions, respectively. Niche overlap was highest during the dry season, lowest at the core-area scale, and increased with spatial resolution. These patterns likely reflected the fine-scale, seasonal habitat preferences of the Cabrera vole, which during the dry season increased the use of small sedge/rush patches embedded in larger tall grass meadows, thereby bringing it in closer contact to the humid habitats selected at coarse spatial resolutions by the water vole throughout the year. Overall, this study suggests that spatial scale may critically influence the perception of habitat differentiation between coexisting species.

Spatial patterns and associations of six congeneric species in an old-growth temperate forest

Analyses of the spatial patterns of pairs of sympatric congeneric species present unique opportunities and challenges in explaining species coexistence. In this study, we compared the population structure and spatial patterns of six coexisting species of Acer (maple) in a large-scale 25-ha old-growth temperate forest plot in northern China with the aim of gaining insight into the coexistence of these congeners. A second-order spatial point pattern analysis based on the pair-correlation function was used to evaluate spatial patterns and examine intra- and interspecies associations among them. The abundance varied from 251 to 6609 individuals, but these species showed similar, distinct L-shaped size-class distributions. All six species were aggregated at small scales. The degree of aggregation of the three species with low abundance was in fact higher than that for the others at small scales. Aggregated patterns were also found for small and large trees at small scales. For spatial associations among the congeners, the goodness-of-fit (GoF) test revealed significant associations for only 10 of 30 species pairs, indicating no clear evidence for interspecific competition within them. In comparing the relationships among different size classes, we found no significant relationship for most of the analyzed species pairs. The GoF test detected significant associations for 6 of 20 species pairs among large trees of different species, 6 of 30 species pairs among small trees of different species, and 11 of 30 species pairs between large and small trees of different species. Based on a comparison of population structure, spatial patterns and associations, we infer that for these congeners there exists no significant difference in seed dispersal, competitive ability, or the requirement for specific topographic and light environments. Other factors (e.g., seed production and other habitat requirements) may contribute to the coexistence of these congeners and the trade-off between species commonness and rarity.