Scale-dependent effects of herbivory on moss communities in Arctic wetlands: A 25-year experiment.

Arctic ecosystems are undergoing rapid changes, including increasing disturbance by herbivore populations, which can affect plant species coexistence and community assemblages. Although the significance of mosses in Arctic wetlands is well recognized, the long-term influence of medium-sized herbivores on the composition of moss communities has received limited attention. We used data from a long-term (25 years) Greater Snow Goose (Anser caerulescens atlanticus) exclusion experiment in Arctic tundra wetlands to assess changes in the composition of moss communities at multiple spatial scales (cell, 4 cm2; quadrat, 100 cm2; exclosure, 16 m2). We investigated how snow goose grazing and grubbing can alter the composition of the moss community by measuring changes in alpha and beta diversity, as well as in the strength of plant interspecific interactions between moss species. Our results indicate that goose foraging significantly increased species diversity (richness, evenness, and inverse Simpson index) of moss communities at the cell and quadrat scales but not the exclosure scale. Goose foraging reduced the dissimilarity (beta diversity) of moss communities at all three scales, mainly due to decreased species turnover. Furthermore, goose foraging increased positive interaction between moss species pairs. These findings emphasize the critical role of geese in promoting moss species coexistence and increasing homogeneity in Arctic wetlands. This study illustrates how top-down regulation by herbivores can alter plant communities in Arctic wetlands and highlights the importance of considering herbivores when examining the response of Arctic plant biodiversity to future climate change.

Effects of different ecological restoration methods on the soil physicochemical properties and vegetation community characteristics of the Baotou light rare earth tailings pond in Inner Mongolia, China.

This study investigated the soil physicochemical properties and vegetation community characteristics of the Baotou light rare earth tailings pond after 10 years of aggregate spray seeding ecological restoration (S1) and ordinary soil spray seeding ecological restoration (S2), and the naturally restored dam slope area without human intervention (S3). The results showed that the vegetation community of S1 was dominated by Caragana korshinskii Kom, and its importance and abundance values were 0.40 and 38.4, respectively, while the vegetation communities of S2 and S3 mainly comprised herbaceous plants. Additionally, the vegetation biomass of S1 was significantly higher than that of S2 and S3 by 215.20% and 1345.76%, respectively, and the vegetation diversity index of S1 was the highest among the three treatment groups. The soil porosity (SP), water content (W), electrical conductivity (EC), and available K were significantly improved in S1, while soil bulk density (BD) was significantly reduced compared with that of S2 and S3. In addition, redundancy analysis revealed that SP, EC, W, and K positively correlate with the biomass, Shannon, Pielou, Simpson, and Marglef indices. Principal component analysis further showed that the comprehensive score of S1 (0.983) was higher than that of S2 (- 0.261) and S3 (- 0.648). Collectively, these findings indicate that appropriate ecological restoration can improve soil structure and vegetation community characteristics, thereby accelerating vegetation restoration, ultimately increasing the stability of the ecosystem.

Adult conspecific density affects Janzen-Connell patterns by modulating the recruitment exclusion zones.

Plant-soil negative feedback (NF) is a well-established phenomenon that, by preventing the dominance of a single species, allows species coexistence and promotes the maintenance of biodiversity. At community scale, localized NF may cause the formation of exclusion zones under adult conspecifics leading to Janzen-Connell (JC) distribution. In this study, we explore the connection between adult density, either conspecifics or heterospecifics, on the probability of occurrence of JC distributions. Using an individual-based modelling approach, we simulated the formation of exclusion zones due to the build-up of NF in proximity of conspecific adult plants and assessed the frequency of JC distribution in relation to conspecifics and heterospecifics density ranging from isolated trees to closed forest stands. We found that JC recruitment distribution is very common in the case of an isolated tree when NF was strong and capable to form an exclusion zone under the parent tree. At very low NF intensity, a prevalence of the decreasing pattern was observed because, under such conditions, the inhibitory effect due to the presence of the mother tree was unable to overcome the clustering effect of the seed dispersal kernel. However, if NF is strong the JC frequency suddenly decreases in stands with a continuous conspecific cover likely as a result of progressive expansion of the exclusion zone surrounding all trees in closed forest stands. Finally, our simulations showed that JC distribution should not be frequent in the case of rare species immersed in a matrix of heterospecific adults. Overall, the model shows that a plant suffering from strong NF in monospecific stands can rarely exhibit a recruitment pattern fitting the JC model. Such counterintuitive results would provide the means to reconcile the well-established NF framework with part the forest ecologists' community that is still skeptical towards the JC model. Synthesis: Our model highlights the complex interconnection between NF intensity, stand density, and recruitment patterns explaining where and why the JC distribution occurs. Moreover, predicting the occurrence of JC in relation to stand density we clarify the relevance of this ecological phenomenon for future integration in plant community frameworks.

Inference of future bog succession trajectory from spatial chronosequence of changing aapa mires.

Climate change-driven vegetation changes can alter the ecosystem functions of northern peatlands. Several case studies have documented fen-to-bog transition (FBT) over recent decades, which can have major implications, as increased bog growth would likely cause cooling feedback. However, studies beyond individual cases are missing to infer if a common trajectory or many alternatives of FBT are in progress. We explored plant community and hydrology patterns during FBT of 23 boreal aapa mire complexes in Finland. We focused on mires where comparisons of historical (1940-1970) and new (2017-2019) aerial photographs indicated an expansion of Sphagnum-dominated zones. Vegetation plot and water chemistry data were collected from string-flark fens, transition zones with indications of Sphagnum increase, and bog zones; thus, in a chronosequence with a decadal time span. We ask, is there a common trajectory or many alternatives of FBT in progress, and what are the main characteristics (species and traits) of transitional plant communities? We found a pattern of fen-bog transitions via an increase in Sphagnum sect. Cuspidata (mainly S. majus and S. balticum), indicating a consistently high water table. Indicators only of transitional communities were scarce (Sphagnum lindbergii), but FBT had apparently facilitated shallow-rooted aerenchymatous vascular plants, especially Scheuchzeria palustris. Water pH consistently reflected the chronosequence with averages of 4.2, 3.9, and 3.8, from fen to transition and bog zones. Due to weak minerotrophy of string-flark fens, species richness increased towards bogs, but succession led to reduced beta diversity and homogenization among bog sites. Decadal chronosequence suggested a future fen-bog transition through a wet phase, instead of a drying trend. Transitional poor-fen vegetation was characterized by the abundance of Sphagnum lindbergii, S. majus, and Scheuchzeria palustris. Sphagnum mosses likely benefit from longer growing seasons and consistently wet and acidic conditions of aapa mires.

Tree recruitment is determined by stand structure and shade tolerance with uncertain role of climate and water relations.

Tree regeneration is a key process for long-term forest dynamics, determining changes in species composition and shaping successional trajectories. While tree regeneration is a highly stochastic process, tree regeneration studies often cover narrow environmental gradients only, focusing on specific forest types or species in distinct regions. Thus, the larger-scale effects of temperature, water availability, and stand structure on tree regeneration are poorly understood.We investigated these effects in respect of tree recruitment (in-growth) along wide environmental gradients using forest inventory data from Flanders (Belgium), northwestern Germany, and Switzerland covering more than 40 tree species. We employed generalized linear mixed models to capture the abundance of tree recruitment in response to basal area, stem density, shade casting ability of a forest stand as well as site-specific degree-day sum (temperature), water balance, and plant-available water holding capacity. We grouped tree species to facilitate comparisons between species with different levels of tolerance to shade and drought.Basal area and shade casting ability of the overstory had generally a negative impact on tree recruitment, but the effects differed between levels of shade tolerance of tree recruitment in all study regions. Recruitment rates of very shade-tolerant species were positively affected by shade casting ability. Stem density and summer warmth (degree-day sum) had similar effects on all tree species and successional strategies. Water-related variables revealed a high degree of uncertainty and did not allow for general conclusions. All variables had similar effects independent of the varying diameter thresholds for tree recruitment in the different data sets.Synthesis: Shade tolerance and stand structure are the main drivers of tree recruitment along wide environmental gradients in temperate forests. Higher temperature generally increases tree recruitment rates, but the role of water relations and drought tolerance remains uncertain for tree recruitment on cross-regional scales.

A native annual forb locally excludes a closely related introduced species that co-occurs in oak-savanna habitat remnants.

Despite the ubiquity of introduced species, their long-term impacts on native plant abundance and diversity remain poorly understood. Coexistence theory offers a tool for advancing this understanding by providing a framework to link short-term individual measurements with long-term population dynamics by directly quantifying the niche and average fitness differences between species. We observed that a pair of closely related and functionally similar annual plants with different origins-native Plectritis congesta and introduced Valerianella locusta-co-occur at the community scale but rarely at the local scale of direct interaction. To test whether niche and/or fitness differences preclude local-scale long-term coexistence, we parameterized models of competitor dynamics with results from a controlled outdoor pot experiment, where we manipulated densities of each species. To evaluate the hypothesis that niche and fitness differences exhibit environmental dependency, leading to community-scale coexistence despite local competitive exclusion, we replicated this experiment with a water availability treatment to determine if this key limiting resource alters the long-term prediction. Water availability impacted population vital rates and intensities of intraspecific versus interspecific competition between P. congesta and V. locusta. Despite environmental influence on competition our model predicts that native P. congesta competitively excludes introduced V. locusta in direct competition across water availability conditions because of an absence of stabilizing niche differences combined with a difference in average fitness, although this advantage weakens in drier conditions. Further, field data demonstrated that P. congesta densities have a negative effect on V. locusta seed prediction. We conclude that native P. congesta limits abundances of introduced V. locusta at the direct-interaction scale, and we posit that V. locusta may rely on spatially dependent coexistence mechanisms to maintain coexistence at the site scale. In quantifying this competitive outcome our study demonstrates mechanistically how a native species may limit the abundance of an introduced invader.

Janzen-Connell effects in a forest BEF experiment: Strong distance-dependent seedling establishment of multiple species.

The Janzen-Connell (JC) hypothesis is a major ecological explanation for high species richness, in particular in tropical forest ecosystems. Central components of the JC hypothesis are noncompetitive effects of distance and density dependence, two drivers that contribute independently to species coexistence, but are ultimately linked in the field. However, although numerous studies provide evidence for either distance- or density-dependent effects based on observational data, experimental testing of simultaneous and interactive effects of distance and density has rarely been conducted, especially in a comprehensive multispecies approach. Here, we make use of the forest Biodiversity-Ecosystem Functioning project (BEF) -China to estimate distance- and density-dependent effects in a reciprocal tree seedling transplant experiment of 11 tree species. We deployed 13,490 juveniles of all 11 species in their own (home) and in all foreign monocultures (away), as well as at three different levels of planting density, thereby testing for distance and density effects, respectively. In addition, to quantify the amount to which density effects were brought about by potential additional effects of intraspecific competition, we set up a common garden experiment with different levels of planting density, where an additional "shadow" treatment controlled for effects of canopy shading. Although the "away" and "high-density" treatments significantly impaired the performance and productivity of seedlings, leaf damage and survival was exclusively affected by either the home/away or the density treatment, respectively. Negative density-dependent effects on leaf damage were less pronounced in the "home" treatment, showing that the effects were not additive. In addition, results obtained in the Common Garden Experiment showed that negative effects of high density may be also brought about by intraspecific competition as an alternative density-responsive mechanism and less by true JC effects. Overall, our results provide strong support on a multispecies basis for the influence of host-specific effects already operating in early stages of a forest plantation. However, they also emphasize the need to account appropriately for potential additional density-responsive mechanisms such as intraspecific competition or microenvironmental conditions when addressing the role of JC effects for species coexistence.

Fruit production is influenced by tree size and size-asymmetric crowding in a wet tropical forest.

In tropical forest communities, seedling recruitment can be limited by the number of fruit produced by adults. Fruit production tends to be highly unequal among trees of the same species, which may be due to environmental factors. We observed fruit production for ~2,000 trees of 17 species across 3 years in a wet tropical forest in Costa Rica. Fruit production was modeled as a function of tree size, nutrient availability, and neighborhood crowding. Following model selection, tree size and neighborhood crowding predicted both the probability of reproduction and the number of fruit produced. Nutrient availability only predicted only the probability of reproduction. In all species, larger trees were more likely to be reproductive and produce more fruit. In addition, number of fruit was strongly negatively related to presence of larger neighboring trees in 13 species; presence of all neighboring trees had a weak-to-moderate negative influence on reproductive status in 16 species. Among various metrics of soil nutrient availability, only sum of base cations was positively associated with reproductive status, and for only four species. Synthesis Overall, these results suggest that direct influences on fruit production tend to be mediated through tree size and crowding from neighboring trees, rather than soil nutrients. However, we found variation in the effects of neighbors and nutrients among species; mechanistic studies of allocation to fruit production are needed to explain these differences.

Crown damage and the mortality of tropical trees.

What causes individual tree death in tropical forests remains a major gap in our understanding of the biology of tropical trees and leads to significant uncertainty in predicting global carbon cycle dynamics. We measured individual characteristics (diameter at breast height, wood density, growth rate, crown illumination and crown form) and environmental conditions (soil fertility and habitat suitability) for 26 425 trees ≥ 10 cm diameter at breast height belonging to 416 species in a 52-ha plot in Lambir Hills National Park, Malaysia. We used structural equation models to investigate the relationships among the different factors and tree mortality. Crown form (a proxy for mechanical damage and other stresses) and prior growth were the two most important factors related to mortality. The effect of all variables on mortality (except habitat suitability) was substantially greater than expected by chance. Tree death is the result of interactions between factors, including direct and indirect effects. Crown form/damage and prior growth mediated most of the effect of tree size, wood density, fertility and habitat suitability on mortality. Large-scale assessment of crown form or status may result in improved prediction of individual tree death at the landscape scale.

Intrinsic biotic factors and microsite conditions drive seedling survival in a species with masting reproduction.

Seedling recruitment following a masting event, where more fruits are produced in synchrony and intermittently compared with other species, plays a crucial role in determining species diversity and community structure. Such seedling recruitment can be superabundant, but followed by high mortality shortly thereafter. Differences in biotic factors such as seedling characteristics, competition, and herbivory, and microsite-specific abiotic factors could determine seedling fate in space and time.In a subtropical forest in south China, for 2 years using censuses conducted every 1-2 months, we monitored 40 seed traps and 120, 1 m2 quadrats in five 1-ha plots located from 1,400 to 1,850 m asl for the masting maple species, Acer campbellii subsp. sinense (Pax) P.C.DeJong. We measured biotic-conspecific and heterospecific seedling density, species richness, herbivory, seedling height, and leaf number-and abiotic-canopy openness, slope, and aspect-factors to assess drivers of seedling survival and evaluated A. campbellii subsp. sinense presence in the soil seed bank (SSB).The masting seed dispersal peak and seedling emergence peak occurred between October 2017 and January 2018, and May 2018, respectively. Of 688 selected seedlings, mortality was 92.7% within one year. No seeds were observed in the SSB. Seedling height and leaf number positively affected seedling survival, while seed placement as measured by aspect also showed effects on survival. Conspecific and heterospecific density and herbivory did not show any clear effect. Higher probabilities of seedling survival were found in areas with larger canopy openness (≥12% canopy gap size) and in steeper microsites (≥35°). Synthesis. Masting is mainly studied as a population-level phenomenon from the fruiting tree perspective. Our study of individual seedling fate revealed that intrinsic biotic factors and seed placement were key drivers of survival. Although biotic determinants such as competition from conspecifics or heterospecifics or herbivory did not determine survival, their ubiquitous presence may be an underlying equalizer in community dynamics where seedlings that overcome biotic pressures, if placed at the right microsite, are at better odds at being recruited to the next life history stages.

Do soil biota influence the outcome of novel interactions between plant competitors?

1. Species are shifting their ranges, for example to higher elevations, in response to climate change. Different plant species and soil microbiota will likely shift their ranges at different rates, giving rise to novel communities of plants and soil organisms. However, the ecological consequences of such novel plant-soil interactions are poorly understood. We experimentally simulated scenarios for novel interactions arising between high- and low elevation plants and soil biota following asynchronous climate change range shifts, asking to what extent the ability of plants to coexist depends on the origin of the soil biota. 2. In a greenhouse experiment, we grew pairs of low- (Poa trivialis and Plantago lanceolata) and high- (Poa alpina and Plantago alpina) elevation plant species alone and against a density gradient of con- or heterospecific neighbours. Plants grew on sterilized field soil that was inoculated with a soil community sampled from either low- or high elevation in the western Swiss Alps. We used the experiment to parameterize competition models, from which we predicted the population-level outcomes of competition in the presence of the different soil biota. 3. In the absence of neighbours, three of the four species produced more biomass with the low elevation soil biota. As a result of generally similar responses across plant species, soil biota tended not to affect plant interaction outcomes, with the low elevation species generally predicted to competitively exclude high elevation species irrespective of the soil biota origin. However, the low elevation grass Poa trivialis was only able to invade communities of Poa alpina in the presence of a low elevation soil biota. This suggests that, at least in some cases, the outcome of novel competitive interactions between plants following climate change will depend on whether shifts in the distribution of plant and soil organisms are asynchronous. 4. Synthesis. Our results indicate that the changing soil communities that plants encounter during range expansion can influence plant performance. However, this is only likely to alter expectations for the ability of plants to coexist following climate change if plant species respond differently to the change in the soil community.

Demographic processes limit upward altitudinal range expansion in a threatened tropical palm.

Understanding the factors that determine species' range limits is a key issue in ecology, and is fundamental for biodiversity conservation under widespread global environmental change. Elucidating how altitudinal variation affects demographic processes may provide important clues for understanding the factors limiting current and future species distributions, yet population dynamics at range limits are still poorly understood. Here, we tested the hypothesis that lower abundance at a species' upper altitudinal range limit is related to lower vital rates. We compared the dynamics of two populations of the tropical palm Euterpe edulis, located near and at the edge of its altitudinal limit of distribution in the Brazilian Atlantic Forest. Data from four annual censuses, from 2012 to 2015, were used. We used matrix population models to estimate asymptotic population growth rates and the elasticity values for the vital rates of the two populations of E. edulis. Life table response experiments were used to compare population performance by measuring the contribution of each vital rate to population growth rates. Population growth rates were not significantly different from one in either population, indicating that both populations were stable during the study period. However, the abundance of all ontogenetic stages was lower at the altitudinal range limit, which was related to decreases in some vital rates, especially fecundity. Additionally, there were higher elasticity values for the survival of immatures and reproductive individuals, compared to all other vital rates, in both populations. Synthesis. Our results show that even a small-scale environmental variation near range limits is sufficient to drive changes in the demography of this threatened palm. A minor increase in elevation approaching the limit of altitudinal distribution may reduce environmental suitability and affect population vital rates, thus contributing to setting upper altitudinal range limits for plants.

Functional community structure of African monodominant Gilbertiodendron dewevrei forest influenced by local environmental filtering.

Monodominant patches of forest dominated by Gilbertiodendron dewevrei are commonly found in central African tropical forests, alongside forests with high species diversity. Although these forests are generally found sparsely distributed along rivers, their occurrence is not thought to be (clearly) driven by edaphic conditions but rather by trait combinations of G. dewevrei that aid in achieving monodominance. Functional community structure between these monodominant and mixed forests has, however, not yet been compared. Additionally, little is known about nondominant species in the monodominant forest community. These two topics are addressed in this study. We investigate the functional community structure of 10 one-hectare plots of monodominant and mixed forests in a central region of the Congo basin, in DR Congo. Thirteen leaf and wood traits are measured, covering 95% (basal area weighted) of all species present in the plots, including leaf nutrient contents, leaf isotopic compositions, specific leaf area, wood density, and vessel anatomy. The trait-based assessment of G. dewevrei shows an ensemble of traits related to water use and transport that could be favorable for its location near forest rivers. Moreover, indications have been found for N and P limitations in the monodominant forest, possibly related to ectomycorrhizal associations formed with G. dewevrei. Reduced leaf N and P contents are found at the community level for the monodominant forest and for different nondominant groups, as compared to those in the mixed forest. In summary, this work shows that environmental filtering does prevail in the monodominant G. dewevrei forest, leading to lower functional diversity in this forest type, with the dominant species showing beneficial traits related to its common riverine locations and with reduced soil N and P availability found in this environment, both coregulating the tree community assembly.

Demographic consequences of greater clonal than sexual reproduction in Dicentra canadensis.

Clonality is a widespread life history trait in flowering plants that may be essential for population persistence, especially in environments where sexual reproduction is unpredictable. Frequent clonal reproduction, however, could hinder sexual reproduction by spatially aggregating ramets that compete with seedlings and reduce inter-genet pollination. Nevertheless, the role of clonality in relation to variable sexual reproduction in population dynamics is often overlooked. We combined population matrix models and pollination experiments to compare the demographic contributions of clonal and sexual reproduction in three Dicentra canadensis populations, one in a well-forested landscape and two in isolated forest remnants. We constructed stage-based transition matrices from 3 years of census data to evaluate annual population growth rates, λ. We used loop analysis to evaluate the relative contribution of different reproductive pathways to λ. Despite strong temporal and spatial variation in seed set, populations generally showed stable growth rates. Although we detected some pollen limitation of seed set, manipulative pollination treatments did not affect population growth rates. Clonal reproduction contributed significantly more than sexual reproduction to population growth in the forest remnants. Only at the well-forested site did sexual reproduction contribute as much as clonal reproduction to population growth. Flowering plants were more likely to transition to a smaller size class with reduced reproductive potential in the following year than similarly sized nonflowering plants, suggesting energy trade-offs between sexual and clonal reproduction at the individual level. Seed production had negligible effects on growth and tuber production of individual plants. Our results demonstrate that clonal reproduction is vital for population persistence in a system where sexual reproduction is unpredictable. The bias toward clonality may be driven by low fitness returns for resource investment in sexual reproduction at the individual level. However, chronic failure in sexual reproduction may exacerbate the imbalance between sexual and clonal reproduction and eventually lead to irreversible loss of sex in the population.

Annual dormancy cycles in buried seeds of shrub species: germination ecology of Sideritis serrata (Labiatae).

The germination ecology of Sideritis serrata was investigated in order to improve ex-situ propagation techniques and management of their habitat. Specifically, we analysed: (i) influence of temperature, light conditions and seed age on germination patterns; (ii) phenology of germination; (iii) germinative response of buried seeds to seasonal temperature changes; (iv) temperature requirements for induction and breaking of secondary dormancy; (v) ability to form persistent soil seed banks; and (vi) seed bank dynamics. Freshly matured seeds showed conditional physiological dormancy, germinating at low and cool temperatures but not at high ones (28/14 and 32/18 °C). Germination ability increased with time of dry storage, suggesting the existence of non-deep physiological dormancy. Under unheated shade-house conditions, germination was concentrated in the first autumn. S. serrata seeds buried and exposed to natural seasonal temperature variations in the shade-house, exhibited an annual conditional dormancy/non-dormancy cycle, coming out of conditional dormancy in summer and re-entering it in winter. Non-dormant seeds were clearly induced into dormancy when stratified at 5 or 15/4 °C for 8 weeks. Dormant seeds, stratified at 28/14 or 32/18 °C for 16 weeks, became non-dormant if they were subsequently incubated over a temperature range from 15/4 to 32/18 °C. S. serrata is able to form small persistent soil seed banks. The maximum seed life span in the soil was 4 years, decreasing with burial depth. This is the second report of an annual conditional dormancy/non-dormancy cycle in seeds of shrub species.

Testing predictions of the Janzen-Connell hypothesis: a meta-analysis of experimental evidence for distance- and density-dependent seed and seedling survival.

The Janzen-Connell hypothesis proposes that specialist natural enemies, such as herbivores and pathogens, maintain diversity in plant communities by reducing survival rates of conspecific seeds and seedlings located close to reproductive adults or in areas of high conspecific density. Variation in the strength of distance- and density-dependent effects is hypothesized to explain variation in plant species richness along climatic gradients, with effects predicted to be stronger in the tropics than the temperate zone and in wetter habitats compared to drier habitats.We conducted a comprehensive literature search to identify peer-reviewed experimental studies published in the 40+ years since the hypothesis was first proposed. Using data from these studies, we conducted a meta-analysis to assess the current weight of evidence for the distance and density predictions of the Janzen-Connell hypothesis.Overall, we found significant support for both the distance- and density-dependent predictions. For all studies combined, survival rates were significantly reduced near conspecifics compared to far from conspecifics, and in areas with high densities of conspecifics compared to areas with low conspecific densities. There was no indication that these results were due to publication bias.The strength of distance and density effects varied widely among studies. Contrary to expectations, this variation was unrelated to latitude, and there was no significant effect of study region. However, we did find a trend for stronger distance and density dependence in wetter sites compared to sites with lower annual precipitation. In addition, effects were significantly stronger at the seedling stage compared to the seed stage.Synthesis. Our study provides support for the idea that distance- and density-dependent mortality occurs in plant communities world-wide. Available evidence suggests that natural enemies are frequently the cause of such patterns, consistent with the Janzen-Connell hypothesis, but additional studies are needed to rule out other mechanisms (e.g. intraspecific competition). With the widespread existence of density and distance dependence clearly established, future research should focus on assessing the degree to which these effects permit species coexistence and contribute to the maintenance of diversity in plant communities.

Trade-offs in seedling growth and survival within and across tropical forest microhabitats.

For niche differences to maintain coexistence of sympatric species, each species must grow and/or survive better than each of the others in at least one set of conditions (i.e., performance trade-offs). However, the extent of niche differentiation in tropical forests remains highly debated. We present the first test of performance trade-offs for wild seedlings in a tropical forest. We measured seedling relative growth rate (RGR) and survival of four common native woody species across 18 light, substrate, and topography microhabitats over 2.5 years within Hawaiian montane wet forest, an ideal location due to its low species diversity and strong species habitat associations. All six species pairs exhibited significant performance trade-offs across microhabitats and for RGR versus survival within microhabitats. We also found some evidence of performance equivalence, with species pairs having similar performance in 26% of comparisons across microhabitats. Across species, survival under low light was generally positively associated with RGR under high light. When averaged over all species, topography (slope, aspect, and elevation) explained most of the variation in RGR attributable to microhabitat variables (51-53%) followed by substrate type (35-37%) and light (11-12%). However, the relative effects of microhabitat differed among species and RGR metric (i.e., RGR for height, biomass, or leaf area). These findings indicate that performance trade-offs among species during regeneration are common in low-diversity tropical forest, although other mechanisms may better explain the coexistence of species with small performance differences.