Bénédicte Bachelot

Ontogenetic shifts in trait-mediated mechanisms of plant community assembly

Identifying the processes that maintain highly diverse plant communities remains a central goal in ecology. Species variation in growth and survival rates across ontogeny, represented by tree size classes and life history stage-specific niche partitioning, are potentially important mechanisms for promoting forest diversity. However, the role of ontogeny in mediating competitive dynamics and promoting functional diversity is not well understood, particular in high-diversity systems such as tropical forests. The interaction between interspecific functional trait variation and ontogenetic shifts in competitive dynamics may yield insights into the ecophysiological mechanisms promoting community diversity. We investigated how functional trait (seed size, maximum height, SLA, leaf N, and wood density) associations with growth, survival, and response to competing neighbors differ among seedlings and two size classes of trees in a subtropical rain forest in Puerto Rico. We used a hierarchical Bayes model of diameter growth and survival to infer trait relationships with ontogenetic change in competitive dynamics. Traits were more strongly associated with average growth and survival than with neighborhood interactions, and were highly consistent across ontogeny for most traits. The associations between trait values and tree responses to crowding by neighbors showed significant shifts as trees grew. Large trees exhibited greater growth as the difference in species trait values among neighbors increased, suggesting trait-associated niche partitioning was important for the largest size class. Our results identify potential axes of niche partitioning and performance-equalizing functional trade-offs across ontogeny, promoting species coexistence in this diverse forest community.

Rare species advantage? Richness of damage types due to natural enemies increases with species abundance in a wet tropical forest

Summary 1.The Janzen–Connell hypothesis (JC) is one potential mechanism to explain the maintenance of high alpha diversity of tree species in tropical forests, operating through differential pressure by natural enemies. 2.We proposed that this differing pressure could arise from the richness of damage types due to natural enemies (RDNE). Following a community compensatory trend (CCT), we hypothesized greater RDNE on common species than on rare species. 3.We evaluated this novel interpretation of the JC by assessing damage patterns on leaves, as a proxy for natural enemy species in 44 tree species. We first evaluated which abiotic and biotic factors affect RDNE. Then, we tested whether increasing RDNE leads to an increasing amount of foliar damage. 4.We found that RDNE Was affected by biotic environments: RDNE increased with mean seedling species abundance. RDNE was higher on species occurring near more closely related neighbours. Was not impacted by abiotic factors. Yet, seedlings of shade-tolerant species hosted a higher RDNE than seedlings of shade-intolerant species. Was positively correlated with amount of foliar damage at the species level. 5.Finally, we tested whether RDNE increased seedling mortality risk. We found that Foliar damage, species abundance and RDNE2 increased mortality risk. Richness of damage types due to natural enemies linearly decreased mortality risk more strongly than RDNE2 increased it. Seedling age decreased mortality risk. 6.Synthesis. The richness of damage types due to natural enemies increased with abundance of the host species, suggesting an important role of enemy diversity in the maintenance of tree diversity. Supporting a novel interpretation of the Janzen–Connell hypothesis, we found a greater mortality risk with increasing RDNE2, but not with increasing RDNE. There was a stronger negative linear effect of RDNE on mortality risk. Rare species with low RDNE as well as species with very high RDNE suffered greater mortality than species hosting intermediate RDNE, reinforcing the complexity of the effect of multiple enemies on prey.

The advantage of the extremes: tree seedlings at intermediate abundance in a tropical forest have the highest richness of above‐ground enemies and suffer the most damage

1. Tropical forest tree diversity has been hypothesized to be maintained via the attraction of density responsive and species-specific enemies. Tests of this hypothesis usually assume a linear relationship between enemy pressure (amount of damage and enemy richness) and seedling or tree density. However, enemy pressure is likely to change nonlinearly with local seedling abundance and community scale tree abundance if enemies are characterized by nonlinear functional responses. 2. We examined the abiotic and biotic factors associated with richness of above-ground enemies and foliar damage found in tree seedlings in a tropical forest in Puerto Rico. Rather than identify specific enemies targeting these seedlings, we used damage morphotypes, a paleo-ecological method, to derive a proxy for enemy species richness. 3. We found that the relationships between local and (conspecific seedling density) community scale (conspecific basal area of adult trees) abundance and both richness of above-ground enemies and foliar damage were hump-shaped. Seedlings of tree species existing at intermediate levels of abundance, at both local and community scales, suffered more damage and experienced pressure from a greater diversity of enemies than those existing at high or low densities. 4. We hypothesized that greater damage at intermediate abundance level could arise from a rich mixture of generalist and specialist enemies targeting seedlings of intermediate abundance tree species. Consistent with this hypothesis, we found that generalist enemies were more diverse on species at rare or intermediate abundance relative to common tree species. However, specialist enemies showed no significant trend across tree species abundance at either the local or community scales. 5. Synthesis. Our results suggest that interspecific variation in tree species abundance leads to differences in the magnitude and type of damage tropical tree seedlings suffer. This variation leads to a nonlinear, hump-shaped relationship between species abundance and enemy damage, highlighting fruitful directions for further development of species coexistence theory.

Interactions among mutualism, competition, and predation foster species coexistence in diverse communities

In natural systems, organisms are simultaneously engaged in mutualistic, competitive, and predatory interactions. Theory predicts that species persistence and community stability are feasible when the beneficial effects of mutualisms are balanced by density-dependent negative feedbacks. Enemy-mediated negative feedbacks can foster plant species coexistence in diverse communities, but empirical evidence remains mixed. Disparity between theoretical expectations and empirical results may arise from the effects of mutualistic mycorrhizal fungi. Here, we build a multiprey species/predator model combined with a bidirectional resource exchange system, which simulates mutualistic interactions between plants and fungi. To reach population persistence, (1) the per capita rate of increase of all plant population must exceed the sum of the negative per capita effects of predation, interspecific competition, and costs of mycorrhizal association, and (2) the per capita numerical response of enemies to mycorrhizal plants must exceed the magnitude of the per capita enemy rate of mortality. These conditions reflect the balance between regulation and facilitation in the system. Interactions between plant natural enemies and mycorrhizal fungi lead to shifts in the strength and direction of net mycorrhizal effects on plants over time, with common plant species deriving greater benefits from mycorrhizal associations than rare plant species.

Nitrogen-fixing trees inhibit growth of regenerating Costa Rican rainforests

Significance Regrowing tropical forests are critical for global biodiversity conservation and carbon capture. Nitrogen availability often controls how fast these forests can regrow. Because nitrogen-fixing plants are the primary source of new nitrogen into these forests, one might expect that more nitrogen fixers lead to faster forest regrowth. However, here we show that nitrogen fixers actually slow forest regrowth. Their competitive influence on neighboring trees outweighs any growth enhancement from their nitrogen inputs at this site. Our results call for a more critical evaluation of how nitrogen fixers influence the surrounding forest, especially given the large uncertainty in global climate projections that hinges on the role of nitrogen fixers during tropical forest regeneration. More than half of the world’s tropical forests are currently recovering from human land use, and this regenerating biomass now represents the largest carbon (C)-capturing potential on Earth. How quickly these forests regenerate is now a central concern for both conservation and global climate-modeling efforts. Symbiotic nitrogen-fixing trees are thought to provide much of the nitrogen (N) required to fuel tropical secondary regrowth and therefore to drive the rate of forest regeneration, yet we have a poor understanding of how these N fixers influence the trees around them. Do they promote forest growth, as expected if the new N they fix facilitates neighboring trees? Or do they suppress growth, as expected if competitive inhibition of their neighbors is strong? Using 17 consecutive years of data from tropical rainforest plots in Costa Rica that range from 10 y since abandonment to old-growth forest, we assessed how N fixers influenced the growth of forest stands and the demographic rates of neighboring trees. Surprisingly, we found no evidence that N fixers facilitate biomass regeneration in these forests. At the hectare scale, plots with more N-fixing trees grew slower. At the individual scale, N fixers inhibited their neighbors even more strongly than did nonfixing trees. These results provide strong evidence that N-fixing trees do not always serve the facilitative role to neighboring trees during tropical forest regeneration that is expected given their N inputs into these systems.

Dynamic preferential allocation to arbuscular mycorrhizal fungi explains fungal succession and coexistence

Evidence accumulates about the role of arbuscular mycorrhizal (AM) fungi in shaping plant communities, but little is known about the factors determining the biomass and coexistence of several types of AM fungi in a plant community. Here, using a consumer-resource framework that treats the relationship between plants and fungi as simultaneous, reciprocal exploitation, we investigated what patterns of dynamic preferential plant carbon allocation to empirically-defined fungal types (on-going partner choice) would be optimal for plants, and how these patterns depend on successional dynamics. We found that ruderal AM fungi can dominate under low steady-state nutrient availability, and competitor AM fungi can dominate at higher steady-state nutrient availability; these are conditions characteristic of early and late succession, respectively. We also found that dynamic preferential allocation alone can maintain a diversity of mutualists, suggesting that on-going partner choice is a new coexistence mechanism for mutualists. Our model can therefore explain both mutualist coexistence and successional strategy, providing a powerful tool to derive testable predictions.

Associations among arbuscular mycorrhizal fungi and seedlings are predicted to change with tree successional status

Arbuscular mycorrhizal (AM) fungi in the soil may influence tropical tree dynamics and forest succession. The mechanisms are poorly understood, because the functional characteristics and abundances of tree species and AM fungi are likely to be codependent. We used generalized joint attribute modeling to evaluate if AM fungi are associated with three forest community metrics for a sub-tropical montane forest in Puerto Rico. The metrics chosen to reflect changes during forest succession are the abundance of seedlings of different successional status, the amount of foliar damage on seedlings of different successional status, and community-weighted mean functional trait values (adult specific leaf area [SLA], adult wood density, and seed mass). We used high-throughput DNA sequencing to identify fungal operational taxonomic units (OTUs) in the soil. Model predictions showed that seedlings of mid- and late-successional species had less leaf damage when the 12 most common AM fungi were abundant compared to when these fungi were absent. We also found that seedlings of mid-successional species were predicted to be more abundant when the 12 most common AM fungi were abundant compared to when these fungi were absent. In contrast, early-successional tree seedlings were predicted to be less abundant when the 12 most common AM fungi were abundant compared to when these fungi were absent. Finally, we showed that, among the 12 most common AM fungi, different AM fungi were correlated with functional trait characteristics of early- or late-successional species. Together, these results suggest that early-successional species might not rely as much as mid- and late-successional species on AM fungi, and AM fungi might accelerate forest succession.

Understanding the recruitment response of juvenile Neotropical trees to logging intensity using functional traits

Selective logging remains a widespread practice in tropical forests, yet the long-term effects of timber harvest on juvenile tree (i.e., sapling) recruitment across the hundreds of species occurring in most tropical forests remain difficult to predict. This uncertainty could potentially exacerbate threats to some of the thousands of timber-valuable tree species in the Amazon. Our objective was to determine to what extent long-term responses of tree species regeneration in logged forests can be explained by their functional traits. We integrate functional trait data for 13 leaf, stem, and seed traits from 25 canopy tree species with a range of life histories, such as the pioneer Goupia glabra and the shade-tolerant Iryanthera hostmannii, together with over 30 yr of sapling monitoring in permanent plots spanning a gradient of harvest intensity at the Paracou Forest Disturbance Experiment (PFDE), French Guiana. We anticipated that more intensive logging would increase recruitment of pioneer species with higher specific leaf area, lower wood densities, and smaller seeds, due to the removal of canopy trees. We define a recruitment response metric to compare sapling regeneration to timber harvest intensity across species. Although not statistically significant, sapling recruitment decreased with logging intensity for eight of 23 species and these species tended to have large seeds and dense wood. A generalized linear mixed model fit using specific leaf area, seed mass, and twig density data explained about 45% of the variability in sapling dynamics. Effects of specific leaf area outweighed those of seed mass and wood density in explaining recruitment dynamics of the sapling community in response to increasing logging intensity. The most intense treatment at the PFDE, which includes stand thinning of non-timber-valuable adult trees and poison-girdling for competitive release, showed evidence of shifting community composition in sapling regeneration at the 30-yr mark, toward species with less dense wood, lighter seeds, and higher specific leaf area. Our results indicate that high-intensity logging can have lasting effects on stand regeneration dynamics and that functional traits can help simplify general trends of sapling recruitment for highly diverse logged tropical forests.