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Title: Demographic trade-offs predict tropical forest dynamics
Understanding tropical forest dynamics and planning for their sustainable management require efficient, yet accurate, predictions of the joint dynamics of hundreds of tree species. With increasing information on tropical tree life histories, our predictive understanding is no longer limited by species data but by the ability of existing models to make use of it. Using a demographic forest model, we show that the basal area and compositional changes during forest succession in a neotropical forest can be accurately predicted by representing tropical tree diversity (hundreds of species) with only five functional groups spanning two essential trade-offs—the growth-survival and stature-recruitment trade-offs. This data-driven modeling framework substantially improves our ability to predict consequences of anthropogenic impacts on tropical forests.
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165 to 168
Sponsoring Org:
National Science Foundation
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  1. Abstract Background and Aims

    Understanding shifts in the demographic and functional composition of forests after major natural disturbances has become increasingly relevant given the accelerating rates of climate change and elevated frequency of natural disturbances. Although plant demographic strategies are often described across a slow–fast continuum, severe and frequent disturbance events influencing demographic processes may alter the demographic trade-offs and the functional composition of forests. We examined demographic trade-offs and the shifts in functional traits in a hurricane-disturbed forest using long-term data from the Luquillo Forest Dynamics Plot (LFPD) in Puerto Rico.


    We analysed information on growth, survival, seed rain and seedling recruitment for 30 woody species in the LFDP. In addition, we compiled data on leaf, seed and wood functional traits that capture the main ecological strategies for plants. We used this information to identify the main axes of demographic variation for this forest community and evaluate shifts in community-weighted means for traits from 2000 to 2016.

    Key Results

    The previously identified growth–survival trade-off was not observed. Instead, we identified a fecundity–growth trade-off and an axis representing seedling-to-adult survival. Both axes formed dimensions independent of resprouting ability. Also, changes in tree species composition during the post-hurricane period reflected a directional shiftmore »from seedling and tree communities dominated by acquisitive towards conservative leaf economics traits and large seed mass. Wood specific gravity, however, did not show significant directional changes over time.


    Our study demonstrates that tree demographic strategies coping with frequent storms and hurricane disturbances deviate from strategies typically observed in undisturbed forests, yet the shifts in functional composition still conform to the expected changes from acquisitive to conservative resource-uptake strategies expected over succession. In the face of increased rates of natural and anthropogenic disturbance in tropical regions, our results anticipate shifts in species demographic trade-offs and different functional dimensions.

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  2. Plant ecophysiological trade-offs between different strategies for tolerating stresses are widely theorized to shape forest functional diversity and vulnerability to climate change. However, trade-offs between hydraulic and stomatal regulation during natural droughts remain under-studied, especially in tropical forests. We investigated eleven mature forest canopy trees in central Amazonia during the strong 2015 El Niño. We found greater xylem embolism resistance (P50 = − 3.3 ± 0.8 MPa) and hydraulic safety margin (HSM = 2.12 ± 0.57 MPa) than previously observed in more precipitation-seasonal rainforests of eastern Amazonia and central America. We also discovered that taller trees exhibited lower embolism resistance and greater stomatal sensitivity, a height-structured trade-off between hydraulic resistance and active stomatal regulation. Such active regulation of tree water status, triggered by the onset of stem embolism, acted as a feedback to avoid further increases in embolism, and also explained declines in photosynthesis and transpiration. These results suggest that canopy trees exhibit a conservative hydraulic strategy to endure drought, with trade-offs between investment in xylem to reduce vulnerability to hydraulic failure, and active stomatal regulation to protect against low water potentials. These findings improve our understanding of strategies in tropical forest canopies and contribute to more accurate prediction of drought responses.
  3. Trait-based analyses provide powerful tools for developing a generalizable, physiologically grounded understanding of how forest communities are responding to ongoing environmental changes. Key challenges lie in (1) selecting traits that best characterize the ecological performance of species in the community and (2) determining the degree and importance of intraspecific variability in those traits. Recent studies suggest that globally evident trait correlations (trait dimensions), such as the leaf economic spectrum, may be weak or absent at local scales. Moreover, trait-based analyses that utilize a mean value to represent a species may be misleading. Mean trait values are particularly problematic if species trait value rankings change along environmental gradients, resulting in species trait crossover. To assess how plant traits (1) covary at local spatial scales, (2) vary across the dominant environmental gradients, and (3) can be partitioned within and across taxa, we collected data on 9 traits for 13 tree species spanning the montane temperate—boreal forest ecotones of New York and northern New England. The primary dimension of the trait ordination was the leaf economic spectrum, with trait variability among species largely driven by differences between deciduous angiosperms and evergreen gymnosperms. A second dimension was related to variability in nitrogen to phosphorousmore »levels and stem specific density. Levels of intraspecific trait variability differed considerably among traits, and was related to variation in light, climate, and tree developmental stage. However, trait rankings across species were generally conserved across these gradients and there was little evidence of species crossover. The persistence of the leaf economics spectrum in both temperate and high-elevation conifer forests suggests that ecological strategies of tree species are associated with trade-offs between resource acquisition and tolerance, and may be quantified with relatively few traits. Furthermore, the assumption that species may be represented with a single trait value may be warranted for some trait-based analyses provided traits were measured under similar light levels and climate conditions.« less
  4. 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.

  5. All organisms experience fundamental conflicts between divergent metabolic processes. In plants, a pivotal conflict occurs between allocation to growth, which accelerates resource acquisition, and to defense, which protects existing tissue against herbivory. Trade-offs between growth and defense traits are not universally observed, and a central prediction of plant evolutionary ecology is that context-dependence of these trade-offs contributes to the maintenance of intraspecific variation in defense [Züst and Agrawal,Annu. Rev. Plant Biol., 68, 513–534 (2017)]. This prediction has rarely been tested, however, and the evolutionary consequences of growth–defense trade-offs in different environments are poorly understood, especially in long-lived species [Cipolliniet al.,Annual Plant Reviews(Wiley, 2014), pp. 263–307]. Here we show that intraspecific trait trade-offs, even when fixed across divergent environments, interact with competition to drive natural selection of tree genotypes corresponding to their growth–defense phenotypes. Our results show that a functional trait trade-off, when coupled with environmental variation, causes real-time divergence in the genetic architecture of tree populations in an experimental setting. Specifically, competitive selection for faster growth resulted in dominance by fast-growing tree genotypes that were poorly defended against natural enemies. This outcome is a signature example of eco-evolutionary dynamics: Competitive interactions affected microevolutionary trajectories on a timescale relevant to subsequentmore »ecological interactions [Brunneret al.,Funct. Ecol.33, 7–12 (2019)]. Eco-evolutionary drivers of tree growth and defense are thus critical to stand-level trait variation, which structures communities and ecosystems over expansive spatiotemporal scales.

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