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1. The operation of ‘negative density-dependence' in seedling cohorts in tropical forests is empirically well-established, but only at a phenomenological level that leaves open the question of why seedlings conspecific with an overtopping parent tree experience higher mortality than heterospecifics. The distinction is of theoretical importance because distinct mechanisms are involved. 2. We consider the two most debated possibilities: seedling mortality resulting from classical Lotka-Volterra density-dependence and seedling mortality resulting from the action of biotic agents, as postulated for the Janzen-Connell mechanism. Our study is the first to identify the full spectrum of mortality factors affecting both conspecific and heterospecific members of seedling cohorts. 3. We took advantage of the occurrence of ‘seedling carpets', dense concentrations of seedlings that appear following fruiting events under reproductive individuals of some species. In these ‘carpets', seedlings conspecific with the overtopping parent tree predominate numerically, but heterospecific seedlings are also typically present. Here, we investigated the differential survival of conspecific versus heterospecific seedlings under focal trees of four species: Calatola microcarpa (Icacinaceae), Clarisia racemosa (Moraceae), Matisia cordata (Bombacaceae/Malvaceae) and Sorocea pileata (Moraceae). 4. We show that mortality rates of conspecific seedlings are much higher than those of heterospecific seedlings and that most conspecific mortality (64-100%) resulted from host-restricted arthropod herbivores and/or fungal pathogens, whereas the mortality of heterospecific seedlings resulted from a variety of other causes. 5. Synthesis. Conspecific seedlings died following attack by apparently host-restricted arthropods or fungi and eventually experienced 100% mortality. The results are inconsistent with classical intra- and inter-specific competition and consistent with the actions of distance-responsive and/or density-responsive ‘enemies', as postulated 40 years ago by Janzen and Connell.

We monitored a close-spaced grid of 289 seed traps in 1.44 ha for 8.4 yr in an Amazonian floodplain forest. In a tree community containing hundreds of species, a median of just three to four species of tree seeds falls annually into each 0.5-m² establishment site. The number of seed species reaching a given site increased linearly with time for the duration of the monitoring period, indicating a roughly random arrival of seed species in a given site-year. The number of seed species captured each year over the entire grid ranged from one-third to one-half of the total captured over the 8.4 yr of monitoring, revealing a substantial temporal component of variation in the seed rain. Seed rain at the 0.5-m² scale displayed extreme spatial variability when all potentially viable seeds were tallied, whereas the rain of dispersed seeds was scant, more nearly uniform, and better mixed. Dispersal limitation, defined as failure of seeds to reach establishment sites, is ≥99% per year for a majority of species, explaining why seed augmentation experiments are often successful. Dispersal limitation has been evoked as an explanation for distance-dependent species turnover in tropical tree communities, but that interpretation contrasts with the fact that many Amazonian tree species possess large geographical ranges that extend for hundreds or thousands of kilometers. A better understanding of the processes that bridge the gap between the scales of seedling establishment and the regulation of forest composition will require new methodologies for studying dispersal on scales larger than those yet achieved.

Pathogens are hypothesized to play an important role in the maintenance of tropical forest plant species richness. Notably, species richness may be promoted by incomplete filling of niche space due interactions of host populations with their pathogens. A potentially important group of pathogens are endophytic fungi, which asymptomatically colonize plants and are diverse and abundant in tropical ecosystems. Endophytes may alter competitive abilities of host individuals and improve host fitness under stress, but may also become pathogenic. Little is known of the impacts of endophytes on niche-space filling of their hosts.Here we evaluate how a widespread fungal endophyte infecting a common tropical palm influences its recruitment and survival in natural ecosystems, and whether this impact is modulated by the abiotic environment, potentially constraining host niche-space filling. Iriartea deltoidea dominates many wet lowland Neotropical forests. Diplodia mutila is a common asymptomatic endophyte in mature plants; however, it causes disease in some seedlings. We investigated the effects of light availability on D. mutila disease expression.We found I. deltoidea seedlings to preferentially occur under shady conditions. Correspondingly, we also found that high light triggers endophyte pathogenicity, while low light favors endosymbiotic development, constraining recruitment of endophyte-infested seedlings to shaded understory by reducing seedling survival in direct light. Pathogenicity of D. mutila under high light is proposed to result from light-induced production of H(2)O(2) by the fungus, triggering hypersensitivity, cell death, and tissue necrosis in the palm. This is the first study to demonstrate that endophytes respond to abiotic factors to influence plant distributions in natural ecosystems; and the first to identify light as a factor influencing where an endophyte is placed on the endosymbiont-pathogen continuum. Our findings show that pathogens can indeed constrain niche-space filling of otherwise successful tropical plant species, providing unoccupied niche space for other species.

1. The term ‘dispersal limitation' represents two distinct component processes: the number of seeds produced (fecundity) and the spatial pattern of the seed rain (distribution). We present a quantitative evaluation of these component processes of dispersal limitation for a tropical forest tree community. 2. Using a regularly spaced grid of 289 seed traps (0.5 m² each), we monitored the seed rain into 1.44 ha of upper Amazonian floodplain forest for 6 years whilst concurrently monitoring sapling recruitment in a 0.81-ha subplot centred within the seed-trapping grid. This arrangement allowed us to compare the spatial pattern of seed rain with that of sapling recruitment. 3. We endeavoured to distinguish between undispersed and dispersed seeds by applying a series of criteria to seeds collected in the traps and by removing from certain analyses all seeds that fell under reproductive conspecifics. Gross fecundity of 30 common species that contribute to the advanced regeneration was uniformly low and the rain of dispersed seeds was lower still, being <1.0 m⁻² year⁻¹ in every case. 4. The rain of dispersed seeds with respect to conspecific reproductives closely matched the recruitment of saplings, whereas gross seed rain (all seeds, including undispersed seeds) did not. 5. Synthesis.‘Dispersal limitation' in this faunally intact Amazonian forest is primarily attributable to a scant rain of dispersed seeds, i.e. fecundity limitation, whereas the distribution of dispersed seeds, being random for most species, appears adequate. Evidence from this and earlier research at the same site indicates that the per-capita success of dispersed seeds is many times higher than that of undispersed seeds. Thus, seed dispersal kernels that do not distinguish between dispersed and undispersed seeds are likely to be biologically misleading.

Natural enemies are known to be important in regulating plant populations and contributing to species coexistence (Janzen-Connell effects). The strength of Janzen-Connell effects (both distance- and density-effects) varies across species, but the life history traits that may mediate such a variation are not well understood. This study examined Janzen-Connell effects across the life stages (seed through adult stages) of two sympatric palm species with distinct phenologies and shade tolerances, two traits that may mediate the strength and timing of Janzen-Connell effects. Populations of two common palm species, Attalea phalerata and Astrocaryum murumuru , were studied in Manu National Park, Peru. Seed predation experiments were conducted to assess Janzen-Connell effects at the seed stage. In the post-seed stages, spatial point pattern analyses of the distributions of individuals and biomass were used to infer the strength of distance- and density-effects. Seed predation was both negative distance- and density-dependent consistent with the Janzen-Connell effects. However, only seedling recruitment for asynchronously fruiting Attalea phalerata was depressed near adults while recruitment remained high for synchronously fruiting Astrocaryum murumuru , consistent with weak distance-effects. Negative density-effects were strong in the early stages for shade-intolerant Attalea phalerata but weak or absent in shade-tolerant Astrocaryum murumuru. Distance- and density-effects varied among the life stages of the two palm species in a manner that corresponded to their contrasting phenology and shade tolerance. Generalizing such connections across many species would provide a route to understanding how trait-mediated Janzen-Connell effects scale up to whole communities of species.

Carnivores have long been used as model organisms to examine mechanisms that allow coexistence among ecologically similar species. Interactions between carnivores, including competition and predation, comprise important processes regulating local community structure and diversity. We use data from an intensive camera-trapping monitoring program across eight Neotropical forest sites to describe the patterns of spatiotemporal organization of a guild of five sympatric cat species: jaguar (Panthera onca), puma (Puma concolor), ocelot (Leopardus pardalis), jaguarundi (Herpailurus yagouaroundi) and margay (Leopardus wiedii). For the three largest cat species, we developed multi-stage occupancy models accounting for habitat characteristics (landscape complexity and prey availability) and models accounting for species interactions (occupancy estimates of potential competitor cat species). Patterns of habitat-use were best explained by prey availability, rather than habitat structure or species interactions, with no evidence of negative associations of jaguar on puma and ocelot occupancy or puma on ocelot occupancy. We further explore temporal activity patterns and overlap of all five felid species. We observed a moderate temporal overlap between jaguar, puma and ocelot, with differences in their activity peaks, whereas higher temporal partitioning was observed between jaguarundi and both ocelot and margay. Lastly, we conducted temporal overlap analysis and calculated species activity levels across study sites to explore if shifts in daily activity within species can be explained by varying levels of local competition pressure. Activity patterns of ocelots, jaguarundis and margays were similarly bimodal across sites, but pumas exhibited irregular activity patterns, most likely as a response to jaguar activity. Activity levels were similar among sites and observed differences were unrelated to competition or intraguild killing risk. Our study reveals apparent spatial and temporal partitioning for most of the species pairs analyzed, with prey abundance being more important than species interactions in governing the local occurrence and spatial distribution of Neotropical forest felids.

Extinction rates in the Anthropocene are three orders of magnitude higher than background and disproportionately occur in the tropics, home of half the world's species. Despite global efforts to combat tropical species extinctions, lack of high-quality, objective information on tropical biodiversity has hampered quantitative evaluation of conservation strategies. In particular, the scarcity of population-level monitoring in tropical forests has stymied assessment of biodiversity outcomes, such as the status and trends of animal populations in protected areas. Here, we evaluate occupancy trends for 511 populations of terrestrial mammals and birds, representing 244 species from 15 tropical forest protected areas on three continents. For the first time to our knowledge, we use annual surveys from tropical forests worldwide that employ a standardized camera trapping protocol, and we compute data analytics that correct for imperfect detection. We found that occupancy declined in 22%, increased in 17%, and exhibited no change in 22% of populations during the last 3-8 years, while 39% of populations were detected too infrequently to assess occupancy changes. Despite extensive variability in occupancy trends, these 15 tropical protected areas have not exhibited systematic declines in biodiversity (i.e., occupancy, richness, or evenness) at the community level. Our results differ from reports of widespread biodiversity declines based on aggregated secondary data and expert opinion and suggest less extreme deterioration in tropical forest protected areas. We simultaneously fill an important conservation data gap and demonstrate the value of large-scale monitoring infrastructure and powerful analytics, which can be scaled to incorporate additional sites, ecosystems, and monitoring methods. In an era of catastrophic biodiversity loss, robust indicators produced from standardized monitoring infrastructure are critical to accurately assess population outcomes and identify conservation strategies that can avert biodiversity collapse.

Defaunation is causing declines of large-seeded animal-dispersed trees in tropical forests worldwide, but whether and how these declines will affect carbon storage across this biome is unclear. Here we show, using a pan-tropical data set, that simulated declines of large-seeded animal-dispersed trees have contrasting effects on aboveground carbon stocks across Earth’s tropical forests. In our simulations, African, American and South Asian forests, which have high proportions of animal-dispersed species, consistently show carbon losses (2–12%), but Southeast Asian and Australian forests, where there are more abiotically dispersed species, show little to no carbon losses or marginal gains (±1%). These patterns result primarily from changes in wood volume, and are underlain by consistent relationships in our empirical data (∼2,100 species), wherein, large-seeded animal-dispersed species are larger as adults than small-seeded animal-dispersed species, but are smaller than abiotically dispersed species. Thus, floristic differences and distinct dispersal mode–seed size–adult size combinations can drive contrasting regional responses to defaunation. Defaunation is linked to the decline of tree species that depend on large animals for seed dispersal, but it is unclear if this affects carbon storage. Here the authors show that defaunation effects on carbon storage vary across continents, driven by relationships between seed dispersal strategies and adult tree size.

Tall canopy trees produce many more seeds than do understory treelets, yet, on average, both classes of trees achieve the same lifetime fitness. Using concurrent data on seedfall (8 years) and sapling recruitment (12 years) from a long-established tree plot at the Cocha Cashu Biological Station in Peru, we show that a 40-m canopy tree must produce roughly 13 times the mass of seeds to generate a sapling as a 5-m understory treelet. Mature tree height accounted for 41% of the variance in seed mass per sapling recruit in a simple univariate regression, whereas a multivariate model that included both intrinsic (seed mass, tree height, and dispersal mode) and extrinsic factors (sapling mortality as a surrogate for microsite quality) explained only 31% of the variance in number of seeds per sapling recruit. The multivariate model accounted for less variance because tall trees produce heavier seeds, on average, than treelets. We used \"intact\" (mostly dispersed) seeds to parameterize the response variable so as to reduce, if not eliminate, any contribution of conspecific crowding to the difference in reproductive efficiency between canopy trees and treelets. Accordingly, a test for negative density dependence failed to expose a relationship between density of reproductive trees in the population and reproductive efficiency (seed mass per recruit). We conclude that understory treelets, some of which produce only a dozen seeds a year, gain their per-seed advantage by failing to attract enemies à la Janzen-Connell, either in ecological or evolutionary time.

The conservation of tropical forest carbon stocks offers the opportunity to curb climate change by reducing greenhouse gas emissions from deforestation and simultaneously conserve biodiversity. However, there has been considerable debate about the extent to which carbon stock conservation will provide benefits to biodiversity in part because whether forests that contain high carbon density in their aboveground biomass also contain high animal diversity is unknown. Here, we empirically examined medium to large bodied ground-dwelling mammal and bird (hereafter \"wildlife\") diversity and carbon stock levels within the tropics using camera trap and vegetation data from a pantropical network of sites. Specifically, we tested whether tropical forests that stored more carbon contained higher wildlife species richness, taxonomic diversity, and trait diversity. We found that carbon stocks were not a significant predictor for any of these three measures of diversity, which suggests that benefits for wildlife diversity will not be maximized unless wildlife diversity is explicitly taken into account; prioritizing carbon stocks alone will not necessarily meet biodiversity conservation goals. We recommend conservation planning that considers both objectives because there is the potential for more wildlife diversity and carbon stock conservation to be achieved for the same total budget if both objectives are pursued in tandem rather than independently. Tropical forests with low elevation variability and low tree density supported significantly higher wildlife diversity. These tropical forest characteristics may provide more affordable proxies of wildlife diversity for future multi-objective conservation planning when fine scale data on wildlife are lacking.