skip to main content


This content will become publicly available on April 8, 2025

Title: Tree diversity reduces variability in sapling survival under drought
Abstract

Enhancing tree diversity may be important to fostering resilience to drought‐related climate extremes. So far, little attention has been given to whether tree diversity can increase the survival of trees and reduce its variability in young forest plantations.

We conducted an analysis of seedling and sapling survival from 34 globally distributed tree diversity experiments (363,167 trees, 168 species, 3744 plots, 7 biomes) to answer two questions: (1) Do drought and tree diversity alter the mean and variability in plot‐level tree survival, with higher and less variable survival as diversity increases? and (2) Do species that survive poorly in monocultures survive better in mixtures and do specific functional traits explain monoculture survival?

Tree species richness reduced variability in plot‐level survival, while functional diversity (Rao's Q entropy) increased survival and also reduced its variability. Importantly, the reduction in survival variability became stronger as drought severity increased. We found that species with low survival in monocultures survived comparatively better in mixtures when under drought. Species survival in monoculture was positively associated with drought resistance (indicated by hydraulic traits such as turgor loss point), plant height and conservative resource‐acquisition traits (e.g. low leaf nitrogen concentration and small leaf size).

Synthesis.The findings highlight: (1) The effectiveness of tree diversity for decreasing the variability in seedling and sapling survival under drought; and (2) the importance of drought resistance and associated traits to explain altered tree species survival in response to tree diversity and drought. From an ecological perspective, we recommend mixing be considered to stabilize tree survival, particularly when functionally diverse forests with drought‐resistant species also promote high survival of drought‐sensitive species.

 
more » « less
Award ID(s):
2044406
NSF-PAR ID:
10500244
Author(s) / Creator(s):
 ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  more » ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;  ;   « less
Publisher / Repository:
Wiley-Blackwell
Date Published:
Journal Name:
Journal of Ecology
ISSN:
0022-0477
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    The maintenance of tree diversity has been explained by multiple mechanisms. One of the most thoroughly studied is conspecific negative density dependence, in which specialist plant enemies reduce survivorship of seeds, seedlings or saplings located near adult conspecifics. Although there is much support that conspecific negative density dependence occurs in temperate forests, only a subset of the species investigated thus far exhibit this recruitment pattern. It remains unclear what drives differential susceptibility to conspecifics among tree species. Previous investigators have considered shade tolerance and mycorrhizal type (arbuscular mycorrhizal vs. ectomycorrhizal association) as two traits that might explain differential susceptibility to conspecific negative density dependence.

    Here, we test whether these two plant traits predict susceptibility of tree saplings to conspecific negative density dependence in a temperate hardwood forest using three responses: spatial point patterns of saplings, sapling growth and sapling survival.

    Spatial patterns of saplings indicate that shade tolerant species are less sensitive to conspecifics than shade intolerant species, but show no differences based on mycorrhizal type. Conversely, shade tolerant saplings exhibit reduced growth, but not survival, when located in areas with high conspecific density. We interpret this finding in light of the conservative functional strategies of shade tolerant species, which typically have low leaf nitrogen levels and slower growth to divert resources to tissue defence against enemies. We found an effect of mycorrhizal type interacting with adult conspecific density, where arbuscular mycorrhizal species show a greater reduction in growth than ectomycorrhizal species in areas dense with conspecifics.

    Synthesis. We conclude that the shade tolerance level and the mycorrhizal type of temperate forest saplings may influence how their growth and survival respond to the adult conspecific trees in their neighbourhoods.

     
    more » « less
  2. Abstract

    The role of tree diversity in restored forests and its impact on key ecological processes like growth and resistance to herbivory has become increasingly important. We analyzed height growth and white‐tailed deerOdocoileus virginianusbrowsing damage to saplings of 16 broadleaved tree species in a large‐scale (13 ha) reforestation experiment in Maryland, USA, where we manipulated tree diversity in 70 1,225‐m2plots. After four growing seasons, higher plot‐level tree richness led to increased deer browsing damage (i.e., associational susceptibility). Despite increased deer damage to saplings in mixed plots, tree richness had no overall effect on sapling height growth. However, diversity–height relationships were related to species functional traits. Light demanding species with large leaves and faster growth rates had reduced heights in mixtures, whereas shade‐tolerant, slower‐growing species generally had either increased or unchanged height growth in diverse tree communities, likely related to increased canopy closure in mixtures relative to monocultures. We show that tree diversity can improve growth of late successional species despite exacerbated mammalian herbivore damage. By facilitating the establishment of species with a range of life‐history strategies, increased tree diversity may enhance ecosystem multi‐functionality in the early stages of forest restoration.

     
    more » « less
  3. Yavitt, Joseph B. (Ed.)
    Conspecific negative density dependence (CNDD) promotes tree species diversity by reducing recruitment near conspecific adults due to biotic feedbacks from herbivores, pathogens, or competitors. While this process is well-described in tropical forests, tests of temperate tree species range from strong positive to strong negative density dependence. To explain this, several studies have suggested that tree species traits may help predict the strength and direction of density dependence: for example, ectomycorrhizal-associated tree species typically exhibit either positive or weaker negative conspecific density dependence. More generally, the strength of density dependence may be predictably related to other species-specific ecological attributes such as shade tolerance, or the relative local abundance of a species. To test the strength of density dependence and whether it affects seedling community diversity in a temperate forest, we tracked the survival of seedlings of three ectomycorrhizal-associated species experimentally planted beneath conspecific and heterospecific adults on the Prospect Hill tract of the Harvard Forest, in Massachusetts, USA. Experimental seedling survival was always lower under conspecific adults, which increased seedling community diversity in one of six treatments. We compared these results to evidence of CNDD from observed sapling survival patterns of 28 species over approximately 8 years in an adjacent 35-ha forest plot. We tested whether species-specific estimates of CNDD were associated with mycorrhizal association, shade tolerance, and local abundance. We found evidence of significant, negative conspecific density dependence (CNDD) in 23 of 28 species, and positive conspecific density dependence in two species. Contrary to our expectations, ectomycorrhizal-associated species generally exhibited stronger (e.g., more negative) CNDD than arbuscular mycorrhizal-associated species. CNDD was also stronger in more shade-tolerant species but was not associated with local abundance. Conspecific adult trees often have a negative influence on seedling survival in temperate forests, particularly for tree species with certain traits. Here we found strong experimental and observational evidence that ectomycorrhizal-associating species consistently exhibit CNDD. Moreover, similarities in the relative strength of density dependence from experiments and observations of sapling mortality suggest a mechanistic link between negative effects of conspecific adults on seedling and sapling survival and local tree species distributions. 
    more » « less
  4. Abstract

    Ecologists often invoke interspecific facilitation to help explain positive biodiversity–ecosystem function relationships in plant communities, but seldom test how it occurs. One mechanism through which one species may facilitate another is by ameliorating abiotic stress. Physiological experiments show that a chronic excess of light can cause stress that depresses carbon assimilation. If shading by a plant's neighbours reduces light stress enough, it may facilitate that plant's growth. If light is instead most often a limiting factor for photosynthesis, shading may have an adverse, competitive effect.

    In a temperate tree diversity experiment, we measured stem growth rates and photosynthetic physiology in broadleaf trees across a gradient of light availability imposed by their neighbours. At the extremes, trees experienced nearly full sun (monoculture), or were shaded by nearby fast‐growing conifers (shaded biculture).

    Most species had slower growth rates with larger neighbours, implying a net competitive effect. On the other hand, the two most shade‐tolerant species (Tilia americanaandAcer negundo) and the most shade‐intolerant one (Betula papyrifera) had faster stem growth rates with larger neighbours. The two shade‐tolerant species had the greatest increases in photoinhibition (reduced dark‐acclimatedFv/Fm) across the gradient of increasing light availability, which suggests they are more vulnerable to chronic light stress. While most species had lower carbon assimilation rates in the shaded biculture treatment,T. americanahad rates up to 25% higher.T. americanaalso dropped its leaves 3–4 weeks earlier in monocultures, curtailing its growing season.

    We conclude that although large neighbours can cause light limitation in shade‐intolerant species, they can also increase growth through abiotic stress amelioration in shade‐tolerant species. Finally, in shade‐intolerantB. papyrifera, we find a pattern of stem elongation in trees with larger neighbours, which suggests that a shade avoidance response may account for the apparent positive trend in stem volume.

    Synthesis. Both positive and negative species interactions in our experiment can be explained in large part by the photosynthetic responses of trees to the light environment created by their neighbours. We show that photosynthetic physiology can help explain the species interactions that underlie biodiversity–ecosystem function relationships. The insights that ecologists gain by searching for such physiological mechanisms may help us forecast species interactions under environmental change.

     
    more » « less
  5. Abstract

    Biodiversity and Ecosystem Function analyses aim to explain how individual species and their interactions affect ecosystem function. With this study, we asked in what ways do species interact, are these interactions affected by species planting pattern, and are initial (planted) proportions or previous year (realized) proportions a better reference point for characterizing grassland diversity effects?

    We addressed these questions with experimental communities compiled from a pool of 16 tallgrass prairie species. We planted all species in monocultures and mixtures that varied in their species richness, evenness, and spatial pattern. We recorded species‐specific biomass production over three growing seasons and fitted Diversity‐Interactions (DI) models to annual plot biomass yields.

    In the establishment season, all species interacted equally to form the diversity effect. In years 2 and 3, each species contributed a unique additive coefficient to its interaction with every other species to form the diversity effect. These interactions were affected byHelianthus maximilianiand the species planting pattern. Models based on species planted proportions better‐fit annual plot yield than models based on species previous contributions to plot biomass.

    Outcomes suggest that efforts to plant tallgrass prairies to maximize diversity effects should focus on the specific species present and in what arrangement they are planted. Furthermore, for particularly diverse grasslands, the effort of collecting annual species biomass data may not be necessary when quantifying diversity effects with DI models.

     
    more » « less