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Mortality of corals is increasing due to bleaching, disease and algal overgrowth. In the Caribbean, low rates of coral recruitment contribute to the slow or undetectable rates of recovery in reef ecosystems. Although algae have long been suspected to interfere with coral recruitment, the mechanisms of that interaction remain unclear. We experimentally tested the effects of turf algal abundance on 3 sequential factors important to recruitment of corals: the biophysical delivery of planktonic coral larvae, their propensity to settle, and the availability of microhabitats where they survive. We deployed coral settlement plates inside and outside damselfishStegastesspp. gardens and cages. Damselfish aggression reduced herbivory from fishes, and cages became fouled with turf algae, both locally increasing algal biomass surrounding the plates. This reduced flushing rates in nursery microhabitats on the plate underside, limiting larvae available for settlement. Coral spat settled preferentially on an early successional crustose coralline algaTitanoderma prototypumbut also on or near other coralline algae, biofilms, and calcareous polychaete worm tubes. Post-settlement survival was highest in the fully grazed, lowest algal biomass treatment, and after 27 mo ‘spat’ densities were 73% higher in this treatment. The ‘gauntlet’ refers to the sequence of ecological processes through which corals must survive to recruit. The highest proportion of coral spat successfully running the gauntlet did so under conditions of low algal biomass resulting from increased herbivory. If coral recruitment is heavily controlled at very local scales by this gauntlet, then coral reef managers could improve a reef’s recruitment potential by managing for reduced algal biomass.

Frugivores shape plant communities via seed dispersal of fleshy‐fruited plant species. However, the structural characteristics that frugivores impart to plant communities are little understood. Evaluating how frugivores structure plant communities via the nonproportional use of available fruit resources is critical to understand the functioning of ecosystems where fleshy‐fruited plant species are dominant, such as tropical forests. We performed a seed‐addition field experiment to investigate how frugivorous birds shape the composition and richness of forests during early stages of secondary succession in cleared areas in Puerto Rico. The experiment tested whether the bird‐generated seed rain and the subsequent early successional plant communities were proportional representations of the fleshy‐fruited species that dominated the surrounding community. Experimental treatments consisted of patches with (a) seed‐additions by wild birds attracted to experimental patches with pole perches, (b) manual seed‐additions proportional to fruit abundance at the local scale (≤50 m from experimental plots) and (c) manual seed‐additions proportional to fruit abundance at the landscape scale (entire study site). Birds' seed‐additions differed in composition and abundance to expectations based on fruit availability at local and landscape scales. Treatments with seeds added by birds had the highest species richness in both the seed rain and the emergence stages despite how, on average, the monthly richness in the landscape‐scale treatment was double that of birds and the local‐scale treatment. This phenomenon was explained by the highest heterogeneity from the bird seed‐addition treatment across months, and the lowest seed per capita emergence rates in landscape treatments. Rather than reflecting relative fruit abundance, birds biased seed rain and per capita emergence towards a non‐random mixture of both small‐ and large‐seeded species, resulting in richer and distinct plant communities. Because frugivory and seed dispersal patterns depart from random encounters between frugivores and plants in communities, successional forests are characterized by an overrepresentation of proportionally rare plant species, and decreases in the dominance of many common species. Thus, for regenerating tropical forests, frugivory can function as mechanism that promotes persistence of rare plant species and their coexistence with more abundant plants. A plain language summary is available for this article. Plain Language Summary

Mechanistic models of community assembly state that biotic and abiotic filters constrain species establishment through selection on their functional traits. Predicting this assembly process is hampered because few studies directly incorporate environmental measurements and scale up from species to community level and because the functional traits' significance is environment dependent. We analyzed community assembly by measuring structure, environmental conditions, and species traits of secondary forests in a species-rich tropical system. We found, as hypothesized, that community structure shaped the local environment and that strong relationships existed between this environment and the traits of the most successful species of the regeneration communities. Path and multivariate analyses showed that temperature and leaf traits that regulate it were the most important factors of community differentiation. Comparisons between the trait composition of the forest's regeneration, juvenile, and adult communities showed a consistent community assembly pattern. These results allowed us to identify the major functional traits and environmental factors involved in the assembly of dry-forest communities and demonstrate that environmental filtering is a predictable and fundamental process of community assembly, even in a complex system such as a tropical forest.

1. Savanna ecosystems span a diverse range of climates, edaphic conditions, and disturbance regimes, the complexity of which has stimulated long-standing interest in the mechanisms that maintain tree-grass coexistence. One hypothesis suggests that tree establishment is strongly limited by one or several demographic bottlenecks at early stages of the tree life cycle. A major impediment to testing this hypothesis is the lack of data on the relative strengths of different bottlenecks across key environmental gradients. 2. To identify demographic bottlenecks that limit early tree establishment (0-18 months), we conducted a series of transplant experiments with two savanna trees species (Acacia robusta and Acacia tortilis) across a natural rainfall and soil fertility gradient in the Serengeti ecosystem, Tanzania. We tested the interactive effects of precipitation, herbivory, seed scarification, grass competition, water limitation, and tree species identity on two key life stages: germination and early seedling survival (0-2 months), and juvenile seedling survival (2-18 months). 3. Germination and early seedling survival increased as a function of rainfall, in the absence of herbivores and when seeds were scarified. Juvenile seedling survival, in contrast, decreased with rainfall but increased in the absence of herbivores. Grass removal had the single strongest (positive) effect on juvenile seedling survival of any treatment. Soil moisture monitoring and grass-addition treatments revealed that grasses negatively affected seedlings in ways that were not necessarily linked to soil moisture. 4. A demographic model combining all effects across early life stages showed that the strength of grass competition on juvenile seedling survival was the key factor limiting early tree establishment. While rainfall had an unexpected opposing effect on the two life stages, the net effect of mean annual precipitation on early tree establishment was positive. 5. Synthesis. Successful tree establishment in Serengeti is maximized by a seemingly unlikely sequence of events: (a) scarification of seeds by browsers, (b) heavy rainfall to promote germination, (c) intensive grazing (but absence of browsers), and (d) dry conditions during juvenile seedling growth (>2 months) to reduce competition with grasses. By considering a wide suite of conditions and their interactions, our experimental results are relevant to ongoing debates about savanna vegetation dynamics and structural shifts in tree:grass ratios.

QUESTIONS: Niche differentiation is a central explanation for the co‐existence and distribution patterns of numerous tree species in tropical forests, but functional equivalence leading to neutral dynamics has been proposed as an alternative explanation. This niche vs neutral debate is fuelled by the highly variable results yielded by studies of the association between tree species distributions and environmental factors, where some studies find strong associations but others do not. Here, we ask how differences in sampling scale between studies contribute to this variation. LOCATION: Barro Colorado Island, Panama. METHODS: Using distribution maps of canopy‐statured individuals, we evaluated patterns of habitat association in five tropical tree species on Barro Colorado Island across a wide range of sampling scales (from 50 to 1600 ha). We investigated the scale‐dependency of species clumping patterns (Ripley's K) and the association of species distributions with important environmental variables (forest age, topography and geological formation) using point pattern analyses. RESULTS: Clump size and clump density had high variances within and among spatial scales. Significant habitat associations were found in all five species, with the number of habitat associations generally increasing with the sampling scale. Ignoring dispersal constraints inflated the number of significant habitat associations. CONCLUSIONS: We found that patterns of habitat association (and hence conclusions on the importance of niche vs neutral processes) are strongly affected by the choice of sampling scale and location. Explicit inclusion of the effect of spatial scale is critical for studies of habitat association and the main processes that structure communities of tropical trees.

Woody encroachment is becoming common in tropical savannas. We studied natural seed rain and performed seed addition experiments in a Brazilian savanna that had not been burned for several decades. We found greater abundance of fire-sensitive species in the seed rain, likely contributing to woody encroachment. Flexible fire management policies that allow for natural and prescribed fires may be required to maintain savanna diversity.

Few invaded ecosystems are free from habitat loss and disturbance, leading to uncertainty whether dominant invasive species are driving community change or are passengers along for the environmental ride. The \"driver\" model predicts that invaded communities are highly interactive, with subordinate native species being limited or excluded by competition from the exotic dominants. The \"passenger\" model predicts that invaded communities are primarily structured by noninteractive factors (environmental change, dispersal limitation) that are less constraining on the exotics, which thus dominate. We tested these alternative hypotheses in an invaded, fragmented, and fire-suppressed oak savanna. We examined the impact of two invasive dominant perennial grasses on community structure using a reduction (mowing of aboveground biomass) and removal (weeding of above-and belowground biomass) experiment conducted at different seasons and soil depths. We examined the relative importance of competition vs. dispersal limitation with experimental seed additions. Competition by the dominants limits the abundance and reproduction of many native and exotic species based on their increased performance with removals and mowing. The treatments resulted in increased light availability and bare soil; soil moisture and N were unaffected. Although competition was limiting for some, 36 of 79 species did not respond to the treatments or declined in the absence of grass cover. Seed additions revealed that some subordinates are dispersal limited; competition alone was insufficient to explain their rarity even though it does exacerbate dispersal inefficiencies by lowering reproduction. While the net effects of the dominants were negative, their presence restricted woody plants, facilitated seedling survival with moderate disturbance (i.e., treatments applied in the fall), or was not the primary limiting factor for the occurrence of some species. Finally, the species most functionally distinct from the dominants (forbs, woody plants) responded most significantly to the treatments. This suggests that relative abundance is determined more by trade-offs relating to environmental conditions (long-term fire suppression) than to traits relating to resource capture (which should most impact functionally similar species). This points toward the passenger model as the underlying cause of exotic dominance, although their combined effects (suppressive and facilitative) on community structure are substantial.

The increase in nutrient availability as a consequence of elevated nitrogen (N) deposition is an important component of global environmental change. This is likely to substantially affect the functioning and provisioning of ecosystem services by drylands, where water and N are often limited. We tested mechanisms of chronic N-enrichment-induced plant species loss in a 10-year field experiment with six levels of N addition rate. Our findings on a semi-arid grassland in Inner Mongolia demonstrated that: (i) species richness (SR) declined by 16 per cent even at low levels of additional N (1.75 g N m–2 yr−1), and 50–70% species were excluded from plots which received high N input (10.5–28 g N m−2 yr−1); (ii) the responses of SR and above-ground biomass (AGB) to N were greater in wet years than dry years; (iii) N addition increased the inter-annual variations in AGB, reduced the drought resistance of production and hence diminished ecosystem stability; (iv) the critical threshold for chronic N-enrichment-induced reduction in SR differed between common and rare species, and increased over the time of the experiment owing to the loss of the more sensitive species. These results clearly indicate that both abundance and functional trait-based mechanisms operate simultaneously on N-induced species loss. The low initial abundance and low above-ground competitive ability may be attributable to the loss of rare species. However, shift from below-ground competition to above-ground competition and recruitment limitation are likely to be the key mechanisms for the loss of abundant species, with soil acidification being less important. Our results have important implications for understanding the impacts of N deposition and global climatic change (e.g. change in precipitation regimes) on biodiversity and ecosystem services of the Inner Mongolian grassland and beyond.

The ability of miniscule larvae to control their fate and replenish populations in dynamic marine environments has been a long-running topic of debate of central importance for managing resources and understanding the ecology and evolution of life in the sea. Larvae are considered to be highly susceptible to offshore transport in productive upwelling regions, thereby increasing dispersal, limiting onshore recruitment, and reducing the intensity of community interactions. We show that 45 species of nearshore crustaceans were not transported far offshore in a recruitment-limited region characterized by strong upwelling. To the contrary, 92% of these larvae remained within 6 km from shore in high densities throughout development along two transects sampled four times during the peak upwelling season. Larvae of most species remained nearshore by remaining below a shallow Ekman layer of seaward-flowing surface waters throughout development. Larvae of other species migrated farther offshore by occurring closer to the surface early in development. Postlarvae evidently returned to nearshore adult habitats either by descending to shoreward-flowing upwelled waters or rising to the sea surface where they can be transported shoreward by wind relaxation events or internal waves. Thus wind-driven offshore transport should not limit recruitment, even in strong upwelling regions, and larvae are more likely to recruit closer to natal populations than is widely believed. This study poses a new challenge to determine the true cause and extent of recruitment limitation for a more diverse array of species along upwelling coasts, and thus to further advance our understanding of the connectivity, dynamics, and structure of coastal populations.

1. Savanna tree cover is dynamic due to disturbances such as fire and herbivory. Frequent fires can limit a key demographic transition from sapling to adult height classes in savanna trees. Saplings may be caught in a 'fire trap', wherein individuals repeatedly resprout following fire top-kill events. Saplings only rarely escape the cycle by attaining a fire-resistant height (e.g. taller than the minimum scorch height) during fire-free intervals. 2. Large mammalian herbivores also may trap trees in shorter size classes. Browsing herbivores directly limit sapling height, while grazing herbivores such as cattle facilitate sapling growth indirectly via grass removal. Experimental studies investigating how meso-wildlife, megaherbivores and domestic livestock affect height of resprouts following fire are rare, but necessary for fully understanding how herbivory may reinforce (or counteract) the fire trap. In our study system, interactive fire-herbivore effects on transitions from sapling (<1 m) to adult tree (>1 m) height classes may be further influenced by plant defences, such as symbiotic ants. 3. We used the Kenya Long-term Exclosure Experiment (KLEE) to investigate how post-fire resprout size of a widespread monodominant East African tree, Acacia drepanolobium was influenced by (a) herbivory by different combinations of cattle, meso-wildlife (15-1,000 kg) and megaherbivores (>1,000 kg) and (b) the presence of acacia-ant mutualists that confer tree defences. We sampled height, stem length and ant occupancy of resprouts exposed to different herbivore combinations before and after controlled burns. 4. Resprout height of saplings that were short prior to fire (<1 m) was reduced primarily by meso-wildlife. Negative effects of elephants on post-fire resprout height increased with pre-fire tree size, suggesting that resprouts of the tallest trees (with the greatest potential to escape the fire trap cycle) were preferentially browsed and reduced in height by elephants. There were no significant cattle effects. 5. Synthesis. We provide experimental evidence for two potential pathways through which large herbivores exert control over sapling escape from the fire trap: (a) post-fire meso-wildlife browsing of short (<1 m) resprouts and (b) elephant browsing of the largest size class of resprouts, which would otherwise be most likely to escape the fire trap.