skip to main content


The NSF Public Access Repository (NSF-PAR) system and access will be unavailable from 11:00 PM ET on Thursday, May 23 until 2:00 AM ET on Friday, May 24 due to maintenance. We apologize for the inconvenience.

Title: Density-dependent adult recruitment in a low-density tropical tree

The Janzen–Connell hypothesis is a well-known explanation for why tropical forests have large numbers of tree species. A fundamental prediction of the hypothesis is that the probability of adult recruitment is less in regions of high conspecific adult density, a pattern mediated by density-dependent mortality in juvenile life stages. Although there is strong evidence in many tree species that seeds, seedlings, and saplings suffer conspecific density-dependent mortality, no study has shown that adult tree recruitment is negatively density dependent. Density-dependent adult recruitment is necessary for the Janzen–Connell mechanism to regulate tree populations. Here, we report density-dependent adult recruitment in the population ofHandroanthus guayacan, a wind-dispersed Neotropical canopy tree species. We use data from high-resolution remote sensing to track individual trees with proven capacity to flower in a lowland moist forest landscape in Panama and analyze these data in a Bayesian framework similar to capture–recapture analysis. We independently quantify probabilities of adult tree recruitment and detection and show that adult recruitment is negatively density dependent. The annualized probability of adult recruitment was 3.03% ⋅ year−1. Despite the detection of negative density dependence in adult recruitment, it was insufficient to stabilize the adult population ofH. guayacan, which increased significantly in size over the decade of observation.

more » « less
Author(s) / Creator(s):
Publisher / Repository:
Proceedings of the National Academy of Sciences
Date Published:
Journal Name:
Proceedings of the National Academy of Sciences
Page Range / eLocation ID:
p. 11268-11273
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    Patterns of seed dispersal and seed mortality influence the spatial structure of plant communities and the local coexistence of competing species. Most seeds are dispersed in proximity to the parent tree, where mortality is also expected to be the highest, because of competition with siblings or the attraction of natural enemies. Whereas distance‐dependent mortality in the seed‐to‐seedling transition was often observed in tropical forests, few studies have attempted to estimate the shape of the survival‐distance curves, which determines whether the peak of seedling establishment occurs away from the parent tree (Janzen–Connell pattern) or if the peak attenuates but remains at the parent location (Hubbell pattern). In this study, we inferred the probability density of seed dispersal and two stages of seedling establishment (new recruits, and seedlings 20 cm or taller) with distance for 24 tree species present in the 50‐ha Forest Dynamics Plot of Barro Colorado Island, Panama. Using data from seed traps, seedling survey quadrats, and tree‐census records spanning the 1988–2014 period, we fit hierarchical Bayesian models including parameters for tree fecundity, the shape of the dispersal kernel, and overdispersion of seed or seedling counts. We combined predictions from multiple dispersal kernels to obtain more robust inferences. We find that Hubbell patterns are the most common and Janzen–Connell patterns are very rare among those species; that distance‐dependent mortality may be stronger in the seed stage, in the early recruit stage, or comparable in both; and that species with larger seeds experience less overall mortality and less distance‐dependent mortality. Finally, we describe how this modeling approach could be extended at a community scale to include less abundant species.

    more » « less
  2. Explaining the maintenance of tropical forest diversity under the countervailing forces of drift and competition poses a major challenge to ecological theory. Janzen−Connell effects, in which host-specific natural enemies restrict the recruitment of juveniles near conspecific adults, provide a potential mechanism. Janzen−Connell is strongly supported empirically, but existing theory does not address the stable coexistence of hundreds of species. Here we use high-performance computing and analytical models to demonstrate that tropical forest diversity can be maintained nearly indefinitely in a prolonged state of transient dynamics due to distance-responsive natural enemies. Further, we show that Janzen−Connell effects lead to community regulation of diversity by imposing a diversity-dependent cost to commonness and benefit to rarity. The resulting species−area and rank−abundance relationships are consistent with empirical results. Diversity maintenance over long time spans does not require dispersal from an external metacommunity, speciation, or resource niche partitioning, only a small zone around conspecific adults in which saplings fail to recruit. We conclude that the Janzen−Connell mechanism can explain the maintenance of tropical tree diversity while not precluding the operation of other niche-based mechanisms such as resource partitioning.

    more » « less
  3. Abstract

    Janzen–Connell effects (JCEs), specialised predation of seeds and seedlings near conspecific trees, are hypothesised to maintain species richness. While previous studies show JCEs can maintain high richness relative to neutral communities, recent theoretical work indicates JCEs may weakly inhibit competitive exclusion when species exhibit interspecific fitness variation. However, recent models make somewhat restrictive assumptions about the functional form of specialised predation—that JCEs occur at a fixed rate when offspring are within a fixed distance of a conspecific tree. Using a theoretical model, I show that the functional form of JCEs largely impacts their ability to maintain coexistence. If predation pressure increases additively with adult tree density and decays exponentially with distance, JCEs maintain considerably higher species richness than predicted by recent models. Loosely parameterising the model with data from a Panamanian tree community, I elucidate the conditions under which JCEs are capable of maintaining high species richness.

    more » « less
  4. Abstract

    Feedbacks between plants and their soil microbial communities often drive negative density dependence in rare, tropical tree species, but their importance to common, temperate trees remains unclear. Additionally, whether negative density dependence is driven by natural enemies (e.g., soil pathogens) or by high densities of seedlings has rarely been assessed. Density dependence may also depend on seedling size, as smaller and/or younger seedlings may be more susceptible to mortality agents. We monitored seedlings ofQuercus rubra, a common, canopy‐dominant temperate tree, to investigate how the density of neighboring adults and seedlings influenced their survival over two years. We assessed how the soil microbial community influenced seedling survival by growing seedlings in a glasshouse inoculated with soil collected from beneath conspecific and heterospecific mature trees. In the field, seedling survival was lower in areas with high densities of mature conspecifics but was unrelated to either conspecific or heterospecific seedling density. Smaller seedlings were also more sensitive than larger seedlings to neighboring adult conspecifics. In the glasshouse, seedlings grown with soil from beneath a conspecific adult had a higher mortality rate than seedlings grown with soil from beneath heterospecific adults or sterilized soil, suggesting that soil microbial communities drive the patterns of mortality in the field. These results illustrate the importance of negative density‐dependent feedbacks resulting from the soil microbial community in a common and ecologically important temperate tree species.

    more » « less
  5. Abstract

    Seedling recruitment can be strongly affected by the composition of nearby plant species. At the neighborhood scale (on the order of tens of meters), adult conspecifics can modify soil chemistry and the presence of host microbes (pathogens and mutualists) across their combined canopy area or rooting zones. At local or small spatial scales (on the order of one to few meters), conspecific seed or seedling density can influence the strength of intraspecific light and resource competition and also modify the density‐dependent spread of natural enemies such as pathogens or invertebrate predators. Intrinsic correlation between proximity to adult conspecifics (i.e., recruitment neighborhood) and local seedling density, arising from dispersal, makes it difficult to separate the independent and interactive factors that contribute to recruitment success. Here, we present a field experiment in which we manipulated both the recruitment neighborhood and seedling density to explore how they interact to influence the growth and survival ofDryobalanops aromatica, a dominant ectomycorrhizal tree species in a Bornean tropical rainforest. First, we found that both local seedling density and recruitment neighborhood had effects on performance ofDaromaticaseedlings, though the nature of these impacts varied between growth and survival. Second, we did not find strong evidence that the effect of density on seedling survival is dependent on the presence of conspecific adult trees. However, accumulation of mutualistic fungi beneath conspecifics adults does facilitate establishment ofDaromaticaseedlings. In total, our results suggest that recruitment near adult conspecifics was not associated with a performance cost and may have weakly benefitted recruiting seedlings. Positive effects of conspecifics may be a factor facilitating the regional hyperabundance of this species. Synthesis: Our results provide support for the idea that dominant species in diverse forests may escape the localized recruitment suppression that limits abundance in rarer species.

    more » « less