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As an abundant element in the Earth's crust, sodium plays an unusual role in food webs. Its availability in terrestrial environments is highly variable, but it is nonessential for most plants, yet essential for animals and most decomposers. Accordingly, sodium requirements are important drivers of various animal behavioural patterns and performance levels. To specifically test whether sodium limitation increases cannibalism in a gregarious lepidopteran herbivore, we hydroponically manipulated Helianthus annuus host plants' tissue-sodium concentrations. Gregarious larvae of the bordered patch butterfly, Chlosyne lacinia, cannibalized siblings when plant-tissue sodium concentrations were low in two separate experiments. Although cannibalism was almost non-existent when sodium concentrations were high, individual mortality rates were also high. Sodium concentration in host plants can have pronounced effects on herbivore behaviour, individual-level performance, and population demographics, all of which are important for understanding the ecology and evolution of plant-animal interactions across a heterogeneous phytochemical landscape., (© 2023. The Author(s).)

Some mistletoe species (Loranthaceae) resemble their host plants to a striking degree. Various mechanisms have been proposed for the developmental origins of novel traits that cause mistletoes to appear similar to their hosts, as well as for the adaptive phenotypic evolution of such traits. Calder (1983) proposed a logically flawed group selectionist seed-dispersal hypothesis for mistletoes to resemble their hosts. Calder's (1983) hypothesis does not provide a viable potential explanation for mistletoe resemblance to hosts., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Plant-soil interactions can be important drivers of biological invasions. In particular, the symbiotic relationship between legumes and nitrogen-fixing soil bacteria (i.e. rhizobia) may be influential in invasion success. Legumes, including Australian acacias, have been introduced into novel ranges around the world. Our goal was to examine the acacia-rhizobia symbiosis to determine whether cointroduction of non-native mutualists plays a role in invasiveness of introduced legumes. To determine whether acacias were introduced abroad concurrently with native symbionts, we selected four species introduced to California (two invasive and two noninvasive in the region) and identified rhizobial strains associating with each species in their native and novel ranges. We amplified three genes to examine phylogenetic placement (16S rRNA) and provenance (nifD and nodC) of rhizobia associating with acacias in California and Australia. We found that all Acacia species, regardless of invasive status, are associating with rhizobia of Australian origin in their introduced ranges, indicating that concurrent acacia-rhizobia introductions have occurred for all species tested. Our results suggest that cointroduction of rhizobial symbionts may be involved in the establishment of non-native acacias in their introduced ranges, but do not contribute to the differential invasiveness of Acacia species introduced abroad., (© The Author(s) 2022. Published by Oxford University Press on behalf of FEMS.)

Understanding the phytochemical landscapes of essential and nonessential chemical elements to plants provides an opportunity to better link biogeochemical cycles to trophic ecology. We investigated the formation and regulation of the cationic phytochemical landscapes of four key elements for biota: Ca, Mg, K, and Na. We collected aboveground tissues of plants in Atriplex , Helianthus , and Opuntia and adjacent soils from 51, 131, and 83 sites, respectively, across the southern United States. We determined the spatial variability of these cations in plants and soils. Also, we quantified the homeostasis coefficient for each cation and genus combination, by using mixed-effect models, with spatially correlated random effects. Additionally, using random forest models, we modeled the influence of bioclimatic, soil, and spatial variables on plant cationic concentrations. Sodium variability and spatial autocorrelation were considerably greater than for Ca, Mg, or K. Calcium, Mg, and K exhibited strongly homeostatic patterns, in striking contrast to non-homeostatic Na. Even so, climatic and soil variables explained a large proportion of plants' cationic concentrations. Essential elements (Ca, Mg, and K) appeared to be homeostatically regulated, which contrasted sharply with Na, a nonessential element for most plants. In addition, we provide evidence for the No-Escape-from-Sodium hypothesis in real-world ecosystems, indicating that plant Na concentrations tend to increase as substrate Na levels increase., Competing Interests: The authors declare no conflict of interest., (© 2022 The Authors. Plant‐Environment Interactions published by New Phytologist Foundation and John Wiley & Sons Ltd.)

As an essential micronutrient for many organisms, sodium plays an important role in ecological and evolutionary dynamics. Although plants mediate trophic fluxes of sodium, from substrates to higher trophic levels, relatively little comparative research has been published about plant growth and sodium accumulation in response to variation in substrate sodium. Accordingly, we carried out a systematic review of plants' responses to variation in substrate sodium concentrations.We compared biomass and tissue-sodium accumulation among 107 cultivars or populations (67 species in 20 plant families), broadly expanding beyond the agricultural and model taxa for which several generalizations previously had been made. We hypothesized a priori response models for each population's growth and sodium accumulation as a function of increasing substrate NaCl and used Bayesian Information Criterion to choose the best model. Additionally, using a phylogenetic signal analysis, we tested for phylogenetic patterning of responses across taxa.The influence of substrate sodium on growth differed across taxa, with most populations experiencing detrimental effects at high concentrations. Irrespective of growth responses, tissue sodium concentrations for most taxa increased as sodium concentration in the substrate increased. We found no strong associations between the type of growth response and the type of sodium accumulation response across taxa. Although experiments often fail to test plants across a sufficiently broad range of substrate salinities, non-crop species tended toward higher sodium tolerance than domesticated species. Moreover, some phylogenetic conservatism was apparent, in that evolutionary history helped predict the distribution of total-plant growth responses across the phylogeny, but not sodium accumulation responses.Our study reveals that saltier plants in saltier soils proves to be a broadly general pattern for sodium across plant taxa. Regardless of growth responses, sodium accumulation mostly followed an increasing trend as substrate sodium levels increased., Competing Interests: The authors have no competing interest to declare., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

When Darwin visited the Galapagos archipelago, he observed that, in spite of the islands' physical similarity, members of species that had dispersed to them recently were beginning to diverge from each other. He postulated that these divergences must have resulted primarily from interactions with sets of other species that had also diverged across these otherwise similar islands. By extrapolation, if Darwin is correct, such complex interactions must be driving species divergences across all ecosystems. However, many current general ecological theories that predict observed distributions of species in ecosystems do not take the details of between-species interactions into account. Here we quantify, in sixteen forest diversity plots (FDPs) worldwide, highly significant negative density-dependent (NDD) components of both conspecific and heterospecific between-tree interactions that affect the trees' distributions, growth, recruitment, and mortality. These interactions decline smoothly in significance with increasing physical distance between trees. They also tend to decline in significance with increasing phylogenetic distance between the trees, but each FDP exhibits its own unique pattern of exceptions to this overall decline. Unique patterns of between-species interactions in ecosystems, of the general type that Darwin postulated, are likely to have contributed to the exceptions. We test the power of our null-model method by using a deliberately modified data set, and show that the method easily identifies the modifications. We examine how some of the exceptions, at the Wind River (USA) FDP, reveal new details of a known allelopathic effect of one of the Wind River gymnosperm species. Finally, we explore how similar analyses can be used to investigate details of many types of interactions in these complex ecosystems, and can provide clues to the evolution of these interactions., Competing Interests: The authors have declared that no competing interests exist. Authors Perry Ong, Zhanqing Hao and Sylvester Tan were unable to confirm their authorship contributions. On their behalf, the corresponding author has reported their contributions to the best of their knowledge.

Spatial patterning is a key natural history attribute of sessile organisms that frequently emerges from and dictates potential for interactions among organisms. We tested whether bunchgrasses, the dominant plant functional group in longleaf pine savanna groundcover communities, are nonrandomly patterned by characterizing the spatial dispersion of three bunchgrass species across six sites in Louisiana and Florida. We mapped bunchgrass tussocks of >5.0 cm basal diameter in three [Formula: see text] plots at each site. We modeled tussocks as two-dimensional objects to analyze their spatial relationships while preserving sizes and shapes of individual tussocks. Tussocks were overdispersed (more regularly spaced than random) for all species and sites at the local interaction scale (<0.3 m). This general pattern likely arises from a tussock-centered, distance-dependent mechanism, for example, intertussock competition. Nonrandom spatial patterns of dominant species have implications for community assembly and ecosystem function in tussock-dominated grasslands and savannas, including those characterized by extreme biodiversity.

We present a meta-analysis of plant responses to fertilization experiments conducted in lowland, species-rich, tropical forests. We also update a key result and present the first species-level analyses of tree growth rates for a 15-yr factorial nitrogen (N), phosphorus (P), and potassium (K) experiment conducted in central Panama. The update concerns community-level tree growth rates, which responded significantly to the addition of N and K together after 10 yr of fertilization but not after 15 yr. Our experimental soils are infertile for the region, and species whose regional distributions are strongly associated with low soil P availability dominate the local tree flora. Under these circumstances, we expect muted responses to fertilization, and we predicted species associated with low-P soils would respond most slowly. The data did not support this prediction, species-level tree growth responses to P addition were unrelated to species-level soil P associations. The meta-analysis demonstrated that nutrient limitation is widespread in lowland tropical forests and evaluated two directional hypotheses concerning plant responses to N addition and to P addition. The meta-analysis supported the hypothesis that tree (or biomass) growth rate responses to fertilization are weaker in old growth forests and stronger in secondary forests, where rapid biomass accumulation provides a nutrient sink. The meta-analysis found no support for the long-standing hypothesis that plant responses are stronger for P addition and weaker for N addition. We do not advocate discarding the latter hypothesis. There are only 14 fertilization experiments from lowland, species-rich, tropical forests, 13 of the 14 experiments added nutrients for five or fewer years, and responses vary widely among experiments. Potential fertilization responses should be muted when the species present are well adapted to nutrient-poor soils, as is the case in our experiment, and when pest pressure increases with fertilization, as it does in our experiment. The statistical power and especially the duration of fertilization experiments conducted in old growth, tropical forests might be insufficient to detect the slow, modest growth responses that are to be expected., (© 2018 by the Ecological Society of America.)

The emergent properties of the collection of species in a natural community, such as diversity and the distribution of relative abundances, are influenced by both niche-based and neutral (stochastic) processes. This pluralistic view of the natural world reconciles theory with empirical observations better than does either a strictly niche- or neutrality-based perspective. Even so, rules (or rules of thumb) that govern the relative contributions that niche-based and stochastic processes make as communities assemble remain only vaguely formulated and incompletely tested. For example, the translation of non-random (non-neutral) ecological processes, which differentially sort among species within a community, into species-compositional patterns may occur more influentially within some demographic subsets of organisms than within others. In other words, the relative contributions of niche vs. neutral processes may vary among age-, size-, or stage-classes. For example, non-random patterns of mortality that occur among seedlings in a rain forest, or among newly settled juveniles in communities of sessile marine communities, could be more influential than non-random mortality during later stages in determining overall community diversity. We propose two alternative, mutually compatible, hypotheses to account for different levels of influence from mortality among life-cycle stages toward producing non-random patterns in organismal communities. The Turnover Model simply posits that those demographic classes characterized by faster rates of turnover contribute greater influence in the short-term as sufficient mortality gives rise to non-random changes to the community, as well as over the longer-term as multiple individuals of a given fast-turnover demographic class transition into later classes compared to each individual that ratchets from a slow-turnover starting class into a later class. The Turnover Model should apply to most communities of organisms. The Niche Model, which posits that niche-based processes are more influential in some demographic classes relative to others, may alternatively or additionally apply to communities. We also propose several alternative mechanisms, especially relevant to forest trees, that could cause dynamics consistent with the Niche Model. These mechanisms depend on differences among demographic classes in the extent of demographic variation that individual organisms experience through their trait values or neighborhood conditions., (© 2017 by the Ecological Society of America.)

While the importance of local-scale habitat niches in shaping tree species turnover along environmental gradients in tropical forests is well appreciated, relatively little is known about the influence of phylogenetic signal in species' habitat niches in shaping local community structure. We used detailed maps of the soil resource and topographic variation within eight 24-50 ha tropical forest plots combined with species phylogenies created from the APG III phylogeny to examine how phylogenetic beta diversity (indicating the degree of phylogenetic similarity of two communities) was related to environmental gradients within tropical tree communities. Using distance-based redundancy analysis we found that phylogenetic beta diversity, expressed as either nearest neighbor distance or mean pairwise distance, was significantly related to both soil and topographic variation in all study sites. In general, more phylogenetic beta diversity within a forest plot was explained by environmental variables this was expressed as nearest neighbor distance versus mean pairwise distance (3.0-10.3 % and 0.4-8.8 % of variation explained among plots, respectively), and more variation was explained by soil resource variables than topographic variables using either phylogenetic beta diversity metric. We also found that patterns of phylogenetic beta diversity expressed as nearest neighbor distance were consistent with previously observed patterns of niche similarity among congeneric species pairs in these plots. These results indicate the importance of phylogenetic signal in local habitat niches in shaping the phylogenetic structure of tropical tree communities, especially at the level of close phylogenetic neighbors, where similarity in habitat niches is most strongly preserved.

Identification of mechanisms that allow some species to outcompete others is a fundamental goal in ecology and invasive species management. One useful approach is to examine congeners varying in invasiveness in a comparative framework across native and invaded ranges. Acacia species have been widely introduced outside their native range of Australia, and a subset of these species have become invasive in multiple parts of the world. Within specific regions, the invasive status of these species varies. Our study examined whether a key mechanism in the life history of Acacia species, the legume-rhizobia symbiosis, influences acacia invasiveness on a regional scale. To assess the extent to which species varying in invasiveness correspondingly differ with regard to the diversity of rhizobia they associate with, we grew seven Acacia species ranging in invasiveness in California in multiple soils from both their native (Australia) and introduced (California) ranges. In particular, the aim was to determine whether more invasive species formed symbioses with a wider diversity of rhizobial strains (i.e. are more promiscuous hosts). We measured and compared plant performance, including aboveground biomass, survival, and nodulation response, as well as rhizobial community composition and richness. Host promiscuity did not differ among invasiveness categories. Acacia species that varied in invasiveness differed in aboveground biomass for only one soil and did not differ in survival or nodulation within individual soils. In addition, acacias did not differ in rhizobial richness among invasiveness categories. However, nodulation differed between regions and was generally higher in the native than introduced range. Our results suggest that all Acacia species introduced to California are promiscuous hosts and that host promiscuity per se does not explain the observed differences in invasiveness within this region. Our study also highlights the utility of assessing potential mechanisms of invasion in species' native and introduced ranges., (© The Authors 2016. Published by Oxford University Press on behalf of the Annals of Botany Company.)

Studies of forest dynamics plots (FDPs) have revealed a variety of negative density-dependent (NDD) demographic interactions, especially among conspecific trees. These interactions can affect growth rate, recruitment and mortality, and they play a central role in the maintenance of species diversity in these complex ecosystems. Here we use an equal area annulus (EAA) point-pattern method to comprehensively analyze data from two tropical FDPs, Barro Colorado Island in Panama and Sinharaja in Sri Lanka. We show that these NDD interactions also influence the continued evolutionary diversification of even distantly related tree species in these FDPs. We examine the details of a wide range of these interactions between individual trees and the trees that surround them. All these interactions, and their cumulative effects, are strongest among conspecific focal and surrounding tree species in both FDPs. They diminish in magnitude with increasing phylogenetic distance between heterospecific focal and surrounding trees, but do not disappear or change the pattern of their dependence on size, density, frequency or physical distance even among the most distantly related trees. The phylogenetic persistence of all these effects provides evidence that interactions between tree species that share an ecosystem may continue to promote adaptive divergence even after the species' gene pools have become separated. Adaptive divergence among taxa would operate in stark contrast to an alternative possibility that has previously been suggested, that distantly related species with dispersal-limited distributions and confronted with unpredictable neighbors will tend to converge on common strategies of resource use. In addition, we have also uncovered a positive density-dependent effect: growth rates of large trees are boosted in the presence of a smaller basal area of surrounding trees. We also show that many of the NDD interactions switch sign rapidly as focal trees grow in size, and that their cumulative effect can strongly influence the distributions and species composition of the trees that surround the focal trees during the focal trees' lifetimes., Competing Interests: The authors have declared that no competing interests exist.

A variety of ecological processes influence diversity and species composition in natural communities. Most of these processes, whether abiotic or biotic, differentially filter individuals from birth to death, thereby altering species' relative abundances. Nonrandom outcomes could accrue throughout ontogeny, or the processes that generate them could be particularly influential at certain stages. One long-standing paradigm in tropical forest ecology holds that patterns of relative abundance among mature trees are largely set by processes operating at the earliest life cycle stages. Several studies confirm filtering processes at some stages, but the longevity of large trees makes a rigorous comparison across size classes impossible without long-term demographic data. Here, we use one of the world's longest-running, plot-based forest dynamics projects to compare nonrandom outcomes across stage classes. We considered a cohort of 7,977 individuals in 186 species that were alive in 1971 and monitored in 13 mortality censuses over 42 y to 2013. Nonrandom mortality with respect to species identity occurred more often in the smaller rather than the larger size classes. Furthermore, observed nonrandom mortality in the smaller size classes had a diversifying influence; species richness of the survivors was up to 30% greater than expected in the two smallest size classes, but not greater than expected in the larger size classes. These results highlight the importance of early life cycle stages in tropical forest community dynamics. More generally, they add to an accumulating body of evidence for the importance of early-stage nonrandom outcomes to community structure in marine and terrestrial environments.

The integration of ecology and evolutionary biology requires an understanding of the evolutionary lability in species' ecological niches. For tropical trees, specialization for particular soil resource and topographic conditions is an important part of the habitat niche, influencing the distributions of individual species and overall tree community structure at the local scale. However, little is known about how these habitat niches are related to the evolutionary history of species. We assessed the relationship between taxonomic rank and tree species' soil resource and topographic niches in eight large (24-50 ha) tropical forest dynamics plots. Niche overlap values, indicating the similarity of two species' distributions along soil or topographic axes, were calculated for all pairwise combinations of co-occurring tree species at each study site. Congeneric species pairs often showed greater niche overlap (i.e., more similar niches) than non-congeneric pairs along both soil and topographic axes, though significant effects were found for only five sites based on Mantel tests. No evidence for taxonomic effects was found at the family level. Our results indicate that local habitat niches of trees exhibit varying degrees of phylogenetic signal at different sites, which may have important ramifications for the phylogenetic structure of these communities.

Tropical tree communities are shaped by local-scale habitat heterogeneity in the form of topographic and edaphic variation, but the life-history stage at which habitat associations develop remains poorly understood. This is due, in part, to the fact that previous studies have not accounted for the widely disparate sample sizes (number of stems) that result when trees are divided into size classes. We demonstrate that the observed habitat structuring of a community is directly related to the number of individuals in the community. We then compare the relative importance of habitat heterogeneity to tree community structure for saplings, juveniles and adult trees within seven large (24-50 ha) tropical forest dynamics plots while controlling for sample size. Changes in habitat structuring through tree life stages were small and inconsistent among life stages and study sites. Where found, these differences were an order of magnitude smaller than the findings of previous studies that did not control for sample size. Moreover, community structure and composition were very similar among tree sub-communities of different life stages. We conclude that the structure of these tropical tree communities is established by the time trees are large enough to be included in the census (1 cm diameter at breast height), which indicates that habitat filtering occurs during earlier life stages.

Both habitat filtering and dispersal limitation influence the compositional structure of forest communities, but previous studies examining the relative contributions of these processes with variation partitioning have primarily used topography to represent the influence of the environment. Here, we bring together data on both topography and soil resource variation within eight large (24-50 ha) tropical forest plots, and use variation partitioning to decompose community compositional variation into fractions explained by spatial, soil resource and topographic variables. Both soil resources and topography account for significant and approximately equal variation in tree community composition (9-34% and 5-29%, respectively), and all environmental variables together explain 13-39% of compositional variation within a plot. A large fraction of variation (19-37%) was spatially structured, yet unexplained by the environment, suggesting an important role for dispersal processes and unmeasured environmental variables. For the majority of sites, adding soil resource variables to topography nearly doubled the inferred role of habitat filtering, accounting for variation in compositional structure that would previously have been attributable to dispersal. Our results, illustrated using a new graphical depiction of community structure within these plots, demonstrate the importance of small-scale environmental variation in shaping local community structure in diverse tropical forests around the globe.

Ecological disturbances frequently control the occurrence and patterning of dominant plants in high-diversity communities like C(4) grasslands and savannas. In such ecosystems disturbance-related processes can have important implications for species, and for whole communities when those species are dominant, yet mechanistic understanding of such processes remains fragmentary. Multiple bunchgrass species commonly co-dominate disturbance-dependent and species-rich pine savannas, where small-scale fuel heterogeneity may influence bunchgrass survival and growth following fires. We quantified how fire in locally varying fuel loads influenced dynamics of dominant C(4) bunchgrasses in a species-rich pine savanna in southeastern Louisiana, USA. We focused on two congeneric, co-dominant species (Schizachyrium scoparium and S. tenerum) with similar growth forms, functional traits and reproductive strategies to highlight effects of fuel heterogeneity during fires. In experimental plots with either reduced or increased fuels versus controls with unmanipulated fuels, we compared: 1) bunchgrass damage and 2) mortality from fires; 3) subsequent growth and 4) flowering. Compared to controls, fire with increased fuels caused greater damage, mortality and subsequent flowering, but did not affect post-fire growth. Fire with reduced fuels had no effect on any of the four measures. The two species responded differently to fire with increased fuels--S. scoparium incurred measurably more damage and mortality than S. tenerum. Logistic regression indicated that the larger average size of S. tenerum tussocks made them resistant to more severe burning where fuels were increased. We speculate that locally increased fuel loading may be important in pine savannas for creating colonization sites because where fuels are light or moderate, dominant bunchgrasses persist through fires. Small-scale heterogeneity in fires, and differences in how species tolerate fire may together promote shared local dominance by different bunchgrasses.

Tropical rain forests play a dominant role in global biosphere-atmosphere CO(2) exchange. Although climate and nutrient availability regulate net primary production (NPP) and decomposition in all terrestrial ecosystems, the nature and extent of such controls in tropical forests remain poorly resolved. We conducted a meta-analysis of carbon-nutrient-climate relationships in 113 sites across the tropical forest biome. Our analyses showed that mean annual temperature was the strongest predictor of aboveground NPP (ANPP) across all tropical forests, but this relationship was driven by distinct temperature differences between upland and lowland forests. Within lowland forests (< 1000 m), a regression tree analysis revealed that foliar and soil-based measurements of phosphorus (P) were the only variables that explained a significant proportion of the variation in ANPP, although the relationships were weak. However, foliar P, foliar nitrogen (N), litter decomposition rate (k), soil N and soil respiration were all directly related with total surface (0-10 cm) soil P concentrations. Our analysis provides some evidence that P availability regulates NPP and other ecosystem processes in lowland tropical forests, but more importantly, underscores the need for a series of large-scale nutrient manipulations - especially in lowland forests - to elucidate the most important nutrient interactions and controls., (© 2011 Blackwell Publishing Ltd/CNRS.)

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 x 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1-10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.

Two prominent mechanisms proposed to structure biodiversity are niche-based ecological filtering and chance arrival of propagules from the species pool. Seed arrival is hypothesized to play a particularly strong role in high-diversity plant communities with large potential species pools and many rare species, but few studies have explored how seed arrival and local ecological filters interactively assemble species-rich communities in space and time. We experimentally manipulated seed arrival and multiple ecological filters in high-diversity, herbaceous-dominated groundcover communities in longleaf pine savannas, which contain the highest small-scale species richness in North America (up to > 40 species/m2). We tested three hypotheses: (1) local communities constitute relatively open-membership assemblages, in which increased seed arrival from the species pool strongly increases species richness; (2) ecological filters imposed by local fire intensity and soil moisture influence recruitment and richness of immigrating species; and (3) ecological filters increase similarity in the composition of immigrating species. In a two-year factorial field experiment, we manipulated local fire intensity by increasing pre-fire fuel loads, soil moisture using rain shelters and irrigation, and seed arrival by adding seeds from the local species pool. Seed arrival increased species richness regardless of fire intensity and soil moisture but interacted with both ecological filters to influence community assembly. High-intensity fire decreased richness of resident species, suggesting an important abiotic filter. In contrast, high-intensity fire increased recruitment and richness of immigrating species, presumably by decreasing effects of other ecological filters (competition and resource limitation) in postfire environments. Drought decreased recruitment and richness of immigrating species, whereas wet soil conditions increased recruitment but decreased or had little effect on richness. Moreover, some ecological filters (wet soil conditions and, to a lesser extent, high-intensity fire) increased similarity in the composition of immigrating species, illustrating conditions that influence deterministic community assembly in species-rich communities. Our experiment provides insights into how dispersal-assembly mechanisms may interact with niche-assembly mechanisms in space (spatial variation in disturbance) and time (temporal variation in resource availability) to structure high-diversity communities and can help guide conservation of threatened longleaf pine ecosystems in the face of habitat fragmentation and environmental change.

Pyrogenic plants dominate many fire-prone ecosystems. Their prevalence suggests some advantage to their enhanced flammability, but researchers have had difficulty tying pyrogenicity to individual-level advantages. Based on our review, we propose that enhanced flammability in fire-prone ecosystems should protect the belowground organs and nearby propagules of certain individual plants during fires. We base this hypothesis on five points: (1) organs and propagules by which many fire-adapted plants survive fires are vulnerable to elevated soil temperatures during fires; (2) the degree to which burning plant fuels heat the soil depends mainly on residence times of fires and on fuel location relative to the soil; (3) fires and fire effects are locally heterogeneous, meaning that individual plants can affect local soil heating via their fuels; (4) how a plant burns can thus affect its fitness; and (5) in many cases, natural selection in fire-prone habitats should therefore favor plants that burn rapidly and retain fuels off the ground. We predict an advantage of enhanced flammability for plants whose fuels influence local fire characteristics and whose regenerative tissues or propagules are affected by local variation in fires. Our "pyrogenicity as protection" hypothesis has the potential to apply to a range of life histories. We discuss implications for ecological and evolutionary theory and suggest considerations for testing the hypothesis.

A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 < or = 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (H(max)) are key traits among forest trees. Seed mass, LMA, WD, and H(max) are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r2 < 0.1 for three and r2 < 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for > 80% of the explained variation and, after WD was included, LMA and H(max) made insignificant contributions. Virtually the full range of values of SM, LMA, and H(max) occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.

Theoretical models predict that effects of dispersal on local biodiversity are influenced by the size and composition of the species pool, as well as ecological filters that limit local species membership. We tested these predictions by conducting a meta-analysis of 28 studies encompassing 62 experiments examining effects of propagule supply (seed arrival) on plant species richness under contrasting intensities of ecological filters (owing to disturbance and resource availability). Seed arrival increased local species richness in a wide range of communities (forest, grassland, montane, savanna, wetland), resulting in a positive mean effect size across experiments. Mean effect size was 70% higher in disturbed relative to undisturbed communities, suggesting that disturbance increases recruitment opportunities for immigrating species. In contrast, effect size was not significantly influenced by nutrient or water availability. Among seed-addition experiments, effect size was positively correlated with species and functional diversity within the pool of added seeds (species evenness and seed-size diversity), primarily in disturbed communities. Our analysis provides experimental support for the general hypothesis that species pools and local environmental heterogeneity interactively structure plant communities. We highlight empirical gaps that can be addressed by future experiments and discuss implications for community assembly, species coexistence, and the maintenance of biodiversity.

In high-diversity communities, rare species encounter one another infrequently and therefore may compete more intensely with common species or guilds for limiting space and resources. In addition, rare species may be strongly recruitment limited because of their low abundances. Under these conditions, stochastic dispersal and immigration history can have an important influence on community structure. We tested the hypothesis that local immigration and competition from common, large-stature guilds interact to structure local biodiversity in high-diversity longleaf pine savanna groundcover assemblages (>30 species/m2). In two factorial field experiments, we increased local immigration by adding seeds of 38 mostly rare, small-stature forbs and sedges to plots physically dominated by either a common, large-stature bunchgrass or shrub species and to plots in which competition from these dominant guilds was reduced. We measured species richness and abundance at two spatial scales (0.01 and 0.25 m2) over two years. Immigration increased total species richness and richness of focal seed addition species regardless of levels of competition with bunchgrasses and shrubs, indicating that many rare, small-stature species can recruit in the face of potential competition from dominant guilds. Removal of dominant guilds increased total and focal species richness in shrub-dominated but not bunchgrass-dominated plots. In addition, competition from both dominant guilds had no clear effect on rank-abundance distributions of focal species. Our results suggest a key role for dispersal assembly in structuring local biodiversity in this high-diversity plant community, but the importance of this mechanism depends on the strength of local niche assembly involving competition from some, but not all, dominant guilds.

Though it is recognized that both stochastic and deterministic processes structure all communities, empirical assessments of their relative importance are rare, particularly within any single community. In this paper, we quantify the dynamic effects of dispersal assembly and niche assembly on the seedling layer in a diverse neotropical rain forest. The two theories make divergent predictions regarding the roles of seed arrival and environmental heterogeneity in generating community structure. Put simply, dispersal assembly posits that the stochasticity inherent to seed arrival structures communities, whereas niche assembly suggests that heterogeneity in post-dispersal environmental conditions is more important. We experimentally sowed 15,132 seeds of eight tree species at varying levels of density and diversity. Every six months we censused the seedlings that germinated and assessed the abiotic and biotic conditions of each plot. We assessed the density, diversity, and species composition of three nested subsets of the seedling layer: seedlings germinated from sown seeds, all seedlings germinated between July 2003 and 2004, and all woody seedlings. We partitioned the variance in density and diversity of each subset of the seedling layer into components representing seed-addition treatments and environmental conditions at 6- to 12-month intervals. Seed additions initially explained more variance in the density and diversity than did environmental heterogeneity for seven of eight sown species, but explained little variance in the density or diversity of the entire seedling layer. Species composition was better explained by seed-addition treatments than by environmental heterogeneity for all three subsets and in all time periods. Nevertheless, the variance in community structure explained by seed-addition treatments declined over the two years following germination, presaging shifts in the relative importance of dispersal assembly and niche assembly. Our study quantifies how dispersal assembly and niche assembly may vary among the components of an ecological community and shift dynamically through time.

A central goal of comparative plant ecology is to understand how functional traits vary among species and to what extent this variation has adaptive value. Here we evaluate relationships between four functional traits (seed volume, specific leaf area, wood density, and adult stature) and two demographic attributes (diameter growth and tree mortality) for large trees of 240 tree species from five Neotropical forests. We evaluate how these key functional traits are related to survival and growth and whether similar relationships between traits and demography hold across different tropical forests. There was a tendency for a trade-off between growth and survival across rain forest tree species. Wood density, seed volume, and adult stature were significant predictors of growth and/or mortality. Both growth and mortality rates declined with an increase in wood density. This is consistent with greater construction costs and greater resistance to stem damage for denser wood. Growth and mortality rates also declined as seed volume increased. This is consistent with an adaptive syndrome in which species tolerant of low resource availability (in this case shade-tolerant species) have large seeds to establish successfully and low inherent growth and mortality rates. Growth increased and mortality decreased with an increase in adult stature, because taller species have a greater access to light and longer life spans. Specific leaf area was, surprisingly, only modestly informative for the performance of large trees and had ambiguous relationships with growth and survival. Single traits accounted for 9-55% of the interspecific variation in growth and mortality rates at individual sites. Significant correlations with demographic rates tended to be similar across forests and for phylogenetically independent contrasts as well as for cross-species analyses that treated each species as an independent observation. In combination, the morphological traits explained 41% of the variation in growth rate and 54% of the variation in mortality rate, with wood density being the best predictor of growth and mortality. Relationships between functional traits and demographic rates were statistically similar across a wide range of Neotropical forests. The consistency of these results strongly suggests that tropical rain forest species face similar trade-offs in different sites and converge on similar sets of solutions.

To explore the importance of 12 elements in litter production and decomposition, we fertilized 36 1600 m(2)-plots with combinations of N, P, K, or micronutrients (i.e. B, Ca, Cu, Fe, Mg, Mn, Mo, S, Zn) for 6 years in a lowland Panamanian forest. The 90% of litter falling as leaves and twigs failed to increase with fertilization, but reproductive litter (fruits and flowers) increased by 43% with N. K enhanced cellulose decomposition; one or more micronutrients enhanced leaf-litter decomposition; P enhanced both. Our results suggest tropical forests are a non-Liebig world of multiple nutrient limitations, with at least four elements shaping rates of litterfall and decomposition. Multiple metallomic enzymes and cofactors likely create gradients in the break down of leaf litter. Selection favours individuals that make more propagules, and even in an N-rich forest, N is a non-substitutable resource for reproduction.

Edge effects are major drivers of change in many fragmented landscapes, but are often highly variable in space and time. Here we assess variability in edge effects altering Amazon forest dynamics, plant community composition, invading species, and carbon storage, in the world's largest and longest-running experimental study of habitat fragmentation. Despite detailed knowledge of local landscape conditions, spatial variability in edge effects was only partially foreseeable: relatively predictable effects were caused by the differing proximity of plots to forest edge and varying matrix vegetation, but windstorms generated much random variability. Temporal variability in edge phenomena was also only partially predictable: forest dynamics varied somewhat with fragment age, but also fluctuated markedly over time, evidently because of sporadic droughts and windstorms. Given the acute sensitivity of habitat fragments to local landscape and weather dynamics, we predict that fragments within the same landscape will tend to converge in species composition, whereas those in different landscapes will diverge in composition. This 'landscape-divergence hypothesis', if generally valid, will have key implications for biodiversity-conservation strategies and for understanding the dynamics of fragmented ecosystems.

Protective floral structures may evolve in response to the negative effects of floral herbivores. For example, water calyces--liquid-filled, cup-like structures resulting from the fusion of sepals--may reduce floral herbivory by submerging buds during their development. Our observations of a water-calyx plant, Chrysothemis friedrichsthaliana (Gesneriaceae), revealed that buds were frequently attacked by ovipositing moths (Alucitidae), whose larvae consumed anthers and stigmas before corollas opened. Almost 25% of per-plant flower production was destroyed by alucitid larvae over two seasons, far exceeding the losses to all other floral herbivores combined. Experimental manipulation of water levels in calyces showed that a liquid barrier over buds halved per-flower alucitid egg deposition and subsequent herbivory, relative to buds in calyces without water. Thus, C. friedrichsthaliana's water calyx helps protect buds from a highly detrimental floral herbivore. Our findings support claims that sepal morphology is largely influenced by selection to reduce floral herbivory, and that these pressures can result in novel morphological adaptations.

Background and Aims: When ecologically important plant traits are correlated they may be said to constitute an ecological 'strategy' dimension. Through identifying these dimensions and understanding their inter-relationships we gain insight into why particular trait combinations are favoured over others and into the implications of trait differences among species. Here we investigated relationships among several traits, and thus the strategy dimensions they represented, across 2134 woody species from seven Neotropical forests., Methods: Six traits were studied: specific leaf area (SLA), the average size of leaves, seed and fruit, typical maximum plant height, and wood density (WD). Trait relationships were quantified across species at each individual forest as well as across the dataset as a whole. 'Phylogenetic' analyses were used to test for correlations among evolutionary trait-divergences and to ascertain whether interspecific relationships were biased by strong taxonomic patterning in the traits., Key Results: The interspecific and phylogenetic analyses yielded congruent results. Seed and fruit size were expected, and confirmed, to be tightly related. As expected, plant height was correlated with each of seed and fruit size, albeit weakly. Weak support was found for an expected positive relationship between leaf and fruit size. The prediction that SLA and WD would be negatively correlated was not supported. Otherwise the traits were predicted to be largely unrelated, being representatives of putatively independent strategy dimensions. This was indeed the case, although WD was consistently, negatively related to leaf size., Conclusions: The dimensions represented by SLA, seed/fruit size and leaf size were essentially independent and thus conveyed largely independent information about plant strategies. To a lesser extent the same was true for plant height and WD. Our tentative explanation for negative WD-leaf size relationships, now also known from other habitats, is that the traits are indirectly linked via plant hydraulics.

The importance of niche vs. neutral assembly mechanisms in structuring tropical tree communities remains an important unsettled question in community ecology [Bell G (2005) Ecology 86:1757-1770]. There is ample evidence that species distributions are determined by soils and habitat factors at landscape (<10(4) km(2)) and regional scales. At local scales (<1 km(2)), however, habitat factors and species distributions show comparable spatial aggregation, making it difficult to disentangle the importance of niche and dispersal processes. In this article, we test soil resource-based niche assembly at a local scale, using species and soil nutrient distributions obtained at high spatial resolution in three diverse neotropical forest plots in Colombia (La Planada), Ecuador (Yasuni), and Panama (Barro Colorado Island). Using spatial distribution maps of >0.5 million individual trees of 1,400 species and 10 essential plant nutrients, we used Monte Carlo simulations of species distributions to test plant-soil associations against null expectations based on dispersal assembly. We found that the spatial distributions of 36-51% of tree species at these sites show strong associations to soil nutrient distributions. Neutral dispersal assembly cannot account for these plant-soil associations or the observed niche breadths of these species. These results indicate that belowground resource availability plays an important role in the assembly of tropical tree communities at local scales and provide the basis for future investigations on the mechanisms of resource competition among tropical tree species.

Forest fragmentation is considered a greater threat to vertebrates than to tree communities because individual trees are typically long-lived and require only small areas for survival. Here we show that forest fragmentation provokes surprisingly rapid and profound alterations in Amazonian tree-community composition. Results were derived from a 22-year study of exceptionally diverse tree communities in 40 1-ha plots in fragmented and intact forests, which were sampled repeatedly before and after fragment isolation. Within these plots, trajectories of change in abundance were assessed for 267 genera and 1,162 tree species. Abrupt shifts in floristic composition were driven by sharply accelerated tree mortality and recruitment within approximately 100 m of fragment margins, causing rapid species turnover and population declines or local extinctions of many large-seeded, slow-growing, and old-growth taxa; a striking increase in a smaller set of disturbance-adapted and abiotically dispersed species; and significant shifts in tree size distributions. Even among old-growth trees, species composition in fragments is being restructured substantially, with subcanopy species that rely on animal seed-dispersers and have obligate outbreeding being the most strongly disadvantaged. These diverse changes in tree communities are likely to have wide-ranging impacts on forest architecture, canopy-gap dynamics, plant-animal interactions, and forest carbon storage.

Most ecological hypotheses about species coexistence hinge on species differences, but quantifying trait differences across species in diverse communities is often unfeasible. We examined the variation of demographic traits using a global tropical forest data set covering 4500 species in 10 large-scale tree inventories. With a hierarchical Bayesian approach, we quantified the distribution of mortality and growth rates of all tree species at each site. This allowed us to test the prediction that demographic differences facilitate species richness, as suggested by the theory that a tradeoff between high growth and high survival allows species to coexist. Contrary to the prediction, the most diverse forests had the least demographic variation. Although demographic differences may foster coexistence, they do not explain any of the 16-fold variation in tree species richness observed across the tropics.

An ecological community's species diversity tends to erode through time as a result of stochastic extinction, competitive exclusion, and unstable host-enemy dynamics. This erosion of diversity can be prevented over the short term if recruits are highly diverse as a result of preferential recruitment of rare species or, alternatively, if rare species survive preferentially, which increases diversity as the ages of the individuals increase. Here, we present census data from seven New and Old World tropical forest dynamics plots that all show the latter pattern. Within local areas, the trees that survived were as a group more diverse than those that were recruited or those that died. The larger (and therefore on average older) survivors were more diverse within local areas than the smaller survivors. When species were rare in a local area, they had a higher survival rate than when they were common, resulting in enrichment for rare species and increasing diversity with age and size class in these complex ecosystems.

We present an analysis of the long-term survival of two cohorts of seedlings of the tropical canopy tree Ocotea whitei (Lauraceae) on a 1-ha plot of mature, lowland moist forest on Barro Colorado Island, Panamá. In 1980, we counted an even-aged cohort of seedlings that germinated in 1979, then measured and tagged survivors in 1981. We also measured and tagged a second, smaller cohort of seedlings that germinated in 1981. We followed the subsequent survival of all seedlings through 1985. Seedling mortality was phenotypically, temporally, and spatially non-random. Important correlates of non-random mortality included: (1) seedling size and age, (2) an El Niño drought, and (3) biotic neighborhood. Larger and older seedlings survived better than smaller and younger seedlings, respectively, and the El Niño-related drought of 1982-1983 was associated with elevated mortality rates. Seedling density, which was strongly correlated with the proximity to the nearest conspecific adult, increased mortality. The observed mortality patterns suggest that processes consistent with the Janzen-Connell hypothesis operate during the recruitment phase of O. whitei population dynamics. However, the processes causing the observed density- and distance-dependent mortality may vary with factors such as total seed number, seedling size, and climatic variation, making it difficult to determine whether time-integrated seedling-to-adult spacing mechanisms other than self-thinning operate on a given plant population. After 6 years in the hectare studied, survivors remained densest and most numerous underneath the adult trees. We conclude that only long-term demographic data, collected at a variety of scales on a variety of species, will ultimately answer the question: do Janzen-Connell effects contribute substantially to structuring tropical forests?

Negative density-dependent recruitment of seedlings, that is, seeds of a given species are less likely to become established seedlings if the density of that species is high, has been proposed to be an important mechanism contributing to the extraordinary diversity of tropical tree communities because it can potentially prevent any particular species from usurping all available space, either in close proximity to seed sources or at relatively larger spatial scales. However, density-dependent recruitment does not necessarily enhance community diversity. Furthermore, although density-dependent effects have been found at some life stages in some species, no study has shown that density-dependent recruitment affects community diversity. Here we report the results of observations in a lowland, moist forest in the Republic of Panamá in which the species identities of 386,027 seeds that arrived at 200 seed traps were compared with the species identities of 13,068 seedlings that recruited into adjacent plots over a 4-year period. Across the 200 sites, recruit seedling diversity was significantly higher than seed diversity. Part of this difference was explained by interspecies differences in average recruitment success. Even after accounting for these differences, however, negative density-dependent recruitment contributes significantly to the increase in diversity from seeds to seedling recruits.

Light gap disturbances have been postulated to play a major role in maintaining tree diversity in species-rich tropical forests. This hypothesis was tested in more than 1200 gaps in a tropical forest in Panama over a 13-year period. Gaps increased seedling establishment and sapling densities, but this effect was nonspecific and broad-spectrum, and species richness per stem was identical in gaps and in nongap control sites. Spatial and temporal variation in the gap disturbance regime did not explain variation in species richness. The species composition of gaps was unpredictable even for pioneer tree species. Strong recruitment limitation appears to decouple the gap disturbance regime from control of tree diversity in this tropical forest.