skip to main content


Title: Large‐ and small‐seeded species have contrasting functional neighborhoods in a subtropical forest
Abstract

Forest community composition is the outcome of multiple forces, including those that increase taxonomic and functional divergence and those that promote convergence in traits. The mechanisms underlying these forces may not operate homogenously within communities; individuals of different species are never perfectly mixed, and thus, species tend to be surrounded and interact with different subsets of species. In fact, taxonomic and functional composition of neighborhoods of different focal species can be highly variable. Here, we examine whether mechanisms driving species‐level neighborhoods relate to intrinsic characteristics of focal species such as differences in life‐history and resource‐uptake strategies and in turn relate to species survival. We focus on two key characteristics: (1) seed mass, which defines a dominant axis of life‐history strategies related to stress tolerance, and (2) understory light preferences that sort species from light‐demanding pioneers to shade‐tolerant. We monitored seedling communities over 10 yr in Puerto Rico and calculated neighborhood trait dispersion in species‐level neighborhoods using seven functional traits. We examined whether species‐level characteristics, seed mass and preferred light conditions, influence patterns of functional dispersion in seedling neighborhoods using linear models. Then, we examined how species‐level functional neighborhoods impact seedling survival. We found that small‐ and large‐seeded species diverge in the type of functional neighborhoods they associate with. Large‐seeded species associate with neighbors that are more similar than expected in leaf economic traits, but more different than expected in seed mass and leaf area traits, while the opposite was found for small‐seeded species. This variation in species functional neighborhood was important in determining seedling survival. In sum, our results suggest that divergent and convergent forces do not operate homogenously over entire communities. Their relative role changes in space, and on a species‐by‐species basis, probably with a deterministic foundation linked to traits such as seed mass.

 
more » « less
Award ID(s):
1754668 1754435
NSF-PAR ID:
10459833
Author(s) / Creator(s):
 ;  ;  ;  ;  ;  ;  ;  
Publisher / Repository:
Wiley Blackwell (John Wiley & Sons)
Date Published:
Journal Name:
Ecosphere
Volume:
11
Issue:
1
ISSN:
2150-8925
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  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.

    Methods

    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 shift 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.

    Conclusions

    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.

     
    more » « less
  2. Abstract

    Understanding the mechanisms that promote the coexistence of hundreds of species over small areas in tropical forest remains a challenge. Many tropical tree species are presumed to be functionally equivalent shade tolerant species but exist on a continuum of performance trade‐offs between survival in shade and the ability to quickly grow in sunlight. These trade‐offs can promote coexistence by reducing fitness differences.

    Variation in plant functional traits related to resource acquisition is thought to predict variation in performance among species, perhaps explaining community assembly across habitats with gradients in resource availability. Many studies have found low predictive power, however, when linking trait measurements to species demographic rates.

    Seedlings face different challenges recruiting on the forest floor and may exhibit different traits and/or performance trade‐offs than older individuals face in the eventual adult niche. Seed mass is the typical proxy for seedling success, but species also differ in cotyledon strategy (reserve vs. photosynthetic) or other leaf, stem and root traits. These can cause species with the same average seed mass to have divergent performance in the same habitat.

    We combined long‐term studies of seedling dynamics with functional trait data collected at a standard life‐history stage in three diverse neotropical forests to ask whether variation in coordinated suites of traits predicts variation among species in demographic performance.

    Across hundreds of species in Ecuador, Panama and Puerto Rico, we found seedlings displayed correlated suites of leaf, stem, and root traits, which strongly correlated with seed mass and cotyledon strategy. Variation among species in seedling functional traits, seed mass, and cotyledon strategy were strong predictors of trade‐offs in seedling growth and survival. These results underscore the importance of matching the ontogenetic stage of the trait measurement to the stage of demographic dynamics.

    Our findings highlight the importance of cotyledon strategy in addition to seed mass as a key component of seed and seedling biology in tropical forests because of the contribution of carbon reserves in storage cotyledons to reducing mortality rates and explaining the growth‐survival trade‐off among species.

    Synthesis: With strikingly consistent patterns across three tropical forests, we find strong evidence for the promise of functional traits to provide mechanistic links between seedling form and demographic performance.

     
    more » « less
  3. Abstract Questions

    We asked: (a) whether the strength of conspecific and heterospecific neighborhood crowding effects on focal tree survival and growth vary with neighborhood radii; and (b) if the relative strength of the effect of neighborhood interactions on tree growth and survival varies with neighborhood scale.

    Location

    Luquillo Forest Dynamics Plot, Puerto Rico.

    Methods

    We used tree survival and growth data and included information on species‐mean trait values related to several leaf traits, maximum height, seed mass and wood density. We incorporated a tree neighborhood modeling approach that uses an area around a focal tree with a specified radius, to describe the interactions between a focal tree and its neighbors. We constructed survival and growth models for each functional trait using a Bayesian approach, and varied the size of the radius from 5 m to 30 m, at 5‐m intervals.

    Results

    The results suggested that the estimated effects of conspecific and heterospecific neighbors on tree performance do not vary based on the size of the neighborhood (5–30 m), suggesting that the effects of conspecific and heterospecific neighbors on the performance of a focal tree likely do not vary substantially beyond a neighborhood radius of 5 m in the Luquillo forest. In contrast, the estimated strength of the functional neighborhood (effect of neighbors based on their functional trait values) on tree performance was dependent on the neighborhood range. Our results also suggested that the effects of trait distances and trait hierarchies on tree survival and growth are acting simultaneously and at the same spatial scales.

    Conclusion

    Findings from this study highlight the importance of spatial scale in community assembly processes, and specifically, call for increased attention when selecting the radius that defines the neighborhood around a focal tree as the selected neighborhood radius influences the community patterns discovered, and affects the conclusions about the drivers that control community assembly.

     
    more » « less
  4. Abstract

    Multidimensional trait frameworks are increasingly used to understand plant strategies for growth and survival. However, it is unclear if frameworks developed at a global level can be applied in local communities and how well these frameworks—based largely on plant morphological traits—align with plant physiology and response to stress.

    We tested the ability of an integrated framework of plant form and function to characterise seedling trait variation and drought response among 22 grasses and forbs common in a semi‐arid grassland. We measured above‐ground and below‐ground traits, and survival to explore how drought response is linked to three trait dimensions (resource conservation, microbial collaboration, and plant size) associated with the framework as well as non‐morphological dimensions (e.g. physiological traits) that are under‐represented in global trait frameworks.

    We found support for three globally‐recognised axes representing trade‐offs in strategies associated with tissue investment (leaf nitrogen, leaf mass per area, root tissue density), below‐ground resource uptake (root diameter, specific root length), and size (shoot mass). However, in contrast to global patterns, above‐ground and below‐ground resource conservation gradients were oppositely aligned: root tissue density was positively correlated with leaf N rather than leaf mass per area. This likely reflects different investment strategies of annual and perennial herbaceous species, as fast‐growing annual species invested in lower density roots and less nitrogen‐rich leaves to maximise plant‐level carbon assimilation. Species with longer drought survival minimised water loss through small above‐ground size and low leaf‐level transpiration rates, and drought survival was best predicted by a principal component axis representing plant size.

    Contrary to our expectations, drought survival in seedlings did not align with the conservation or collaboration axes suggesting that seedlings with different functional strategies can achieve similar drought survival, as long as they minimise water loss. Our results also show that within local communities, expected trait relationships could be decoupled as some plant groups achieve similar performance through different trait combinations. The effectiveness of species mean trait values in predicting drought response highlights the value of trait‐based methods as a versatile tool for understanding ecological processes locally across various ecosystems.

    Read the freePlain Language Summaryfor this article on the Journal blog.

     
    more » « less
  5. Disturbance plays a key role in shaping forest composition and diversity. We used a community phylogeny and long-term forest dynamics data to investigate biotic and abiotic factors shaping tropical forest regeneration following both human and natural disturbance. Specifically, we examined shifts in seedling phylogenetic and functional (i.e., seed mass) community structure over a decade following a major hurricane in a human-impacted forest in Puerto Rico. Phylogenetic relatedness of the seedling community decreased in the first five years post-hurricane and then increased, largely driven by changes in the abundance of a common palm species. Functional structure (based on seed mass) became increasingly clustered through time, due to canopy closure causing small-seeded, light-demanding species to decline in abundance. Seedling neighbor density and phylogenetic relatedness negatively affected seedling survival, which likely acted to reduce phylogenetic relatedness within seedling plots. Across the study site, areas impacted in the past by high-intensity land use had lower or similar phylogenetic relatedness of seedling communities than low-intensity past land use areas, reflecting interactive effects of human and natural disturbance. Our study demonstrates how phylogenetic and functional information offer insights into the role of biotic and abiotic factors structuring forest recovery following disturbance. 
    more » « less