skip to main content


Title: Environmental heterogeneity and biotic interactions mediate climate impacts on tropical forest regeneration
Abstract

Predicting the fate of tropical forests under a changing climate requires understanding species responses to climatic variability and extremes. Seedlings may be particularly vulnerable to climatic stress given low stored resources and undeveloped roots; they also portend the potential effects of climate change on future forest composition. Here we use data for ca. 50,000 tropical seedlings representing 25 woody species to assess (i) the effects of interannual variation in rainfall and solar radiation between 2007 and 2016 on seedling survival over 9 years in a subtropical forest; and (ii) how spatial heterogeneity in three environmental factors—soil moisture, understory light, and conspecific neighborhood density—modulate these responses. Community‐wide seedling survival was not sensitive to interannual rainfall variability but interspecific variation in these responses was large, overwhelming the average community response. In contrast, community‐wide responses to solar radiation were predominantly positive. Spatial heterogeneity in soil moisture and conspecific density were the predominant and most consistent drivers of seedling survival, with the majority of species exhibiting greater survival at low conspecific densities and positive or nonlinear responses to soil moisture. This environmental heterogeneity modulated impacts of rainfall and solar radiation. Negative conspecific effects were amplified during rainy years and at dry sites, whereas the positive effects of radiation on survival were more pronounced for seedlings existing at high understory light levels. These results demonstrate that environmental heterogeneity is not only the main driver of seedling survival in this forest but also plays a central role in buffering or exacerbating impacts of climate fluctuations on forest regeneration. Since seedlings represent a key bottleneck in the demographic cycle of trees, efforts to predict the long‐term effects of a changing climate on tropical forests must take into account this environmental heterogeneity and how its effects on regeneration dynamics play out in long‐term stand dynamics.

 
more » « less
Award ID(s):
1754668
NSF-PAR ID:
10048608
Author(s) / Creator(s):
 ;  ;  
Publisher / Repository:
Wiley-Blackwell
Date Published:
Journal Name:
Global Change Biology
Volume:
24
Issue:
2
ISSN:
1354-1013
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    Tropical forest understory regeneration occurs rapidly after disturbance with compositional trajectories that depend on species availability and environmental conditions. To predict future tropical forest regeneration dynamics, we need a deeper understanding of how pulse disturbance events, like hurricanes, interact with environmental variability to affect understory demography and composition. We examined fern and sapling mortality, recruitment, and community composition in relation to solar radiation and soil moisture using 17 years of forest dynamics data (2003–2019) from the Canopy Trimming Experiment in the Luquillo Experimental Forest, Puerto Rico. Solar radiation increased 150% and soil moisture increased 40% following canopy trimming of experimental plots relative to control plots. All plots were disturbed in 2017 by Hurricanes Irma and Maria, so experimentally trimmed plots presented the opportunity to study the effects of multiple hurricanes, while control plots isolated the effects of a single natural hurricane. Recruitment rates maximized at 0.14 individuals/plot/month for ferns and 0.20 stems/plot/month for saplings. Recruitment and mortality were distributed more evenly over the 17 years of monitoring in experimentally trimmed plots than in control plots; however, following Hurricane Maria demographic rates substantially increased in control plots only. In experimentally trimmed plots, the largest community compositional shifts occurred as a result of the trimming events, and compositional changes were greatest for control plots after Hurricane Maria in 2017. Pioneer tree and fern species increased in abundance in response to both simulated and natural hurricanes. Following Hurricane Maria, two dominant pioneer species,Cyathea arboreaandCecropia schreberiana, recruited abundantly, but only in control plots. In trimmed plots, increased solar radiation and soil moisture shifted understory species composition steadily toward pioneer and secondary‐successional species, with soil moisture interacting strongly with canopy trimming. Thus, both solar radiation and soil moisture are environmental drivers affecting pioneer species recruitment following disturbance, which interact with canopy opening following hurricanes. Our results suggest that if hurricane disturbances increase in frequency and severity, as suggested by climate change predictions, the understory regeneration of late‐successional species, such asManilkara bidentataandSloanea berteroana, which prefer deeper shade and slightly drier soil microsites, may become imperiled.

     
    more » « less
  2. Global climate change has led to rising temperatures and to more frequent and intense climatic events, such as storms and droughts. Changes in climate and disturbance regimes can have non-additive effects on plant communities and result in complicated legacies we have yet to understand. This is especially true for tropical forests, which play a significant role in regulating global climate. We used understory vegetation data from the Tropical Responses to Altered Climate Experiment (TRACE) in Puerto Rico to evaluate how plant communities responded to climate warming and disturbance. The TRACE understory vegetation was exposed to a severe drought (2015), 2 years of experimental warming (4°C above ambient in half of the plots, 2016–2017 and 2018–2019), and two major hurricanes (Irma and María, September 2017). Woody seedlings and saplings were censused yearly from 2015 to 2019, with an additional census in 2015 after the drought ended. We evaluated disturbance-driven changes in species richness, diversity, and composition across ontogeny. We then used Bayesian predictive trait modeling to assess how species responded to disturbance and how this might influence the functional structure of the plant community. Our results show decreased seedling richness after hurricane disturbance, as well as increased sapling richness and diversity after warming. We found a shift in species composition through time for both seedlings and saplings, yet the individual effects of each disturbance were not significant. At both ontogenetic stages, we observed about twice as many species responding to experimental warming as those responding to drought and hurricanes. Predicted changes in functional structure point to disturbance-driven functional shifts toward a mixture of fast-growing and drought-tolerant species. Our findings demonstrate that the tropical forest understory community is more resistant to climatic stressors than expected, especially at the sapling stage. However, early signs of changes in species composition suggest that, in a warming climate with frequent droughts and hurricanes, plant communities might shift over time toward fast-growing or drought-tolerant species. 
    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

    Ectomycorrhizal tree species may benefit from positive plant–soil feedbacks, where soil environments near adult trees enhance conspecific seedling growth and survival. In tropical monodominant forests, seedling survival is particularly important, as seedling banks help maintain stand‐level dominance over generations. Positive plant–soil feedbacks may be mediated by diverse ectomycorrhizal fungal communities, which improve nutrient acquisition of heavily shaded seedlings. Despite the potential importance of these fungi, little is known about ectomycorrhizal fungal community development on seedlings of tropical monodominant trees. In Guyana, we sequentially monitored percent colonization and species composition of ectomycorrhizal fungi on an even‐age cohort of seedlings of the tropical monodominant treeDicymbe corymbosa(Fabaceae subfamily Detarioideae). Ectomycorrhizal fungi found onDcorymbosaseedlings over a 12‐month period of early development were compared to those of conspecific adults and four other ectomycorrhizal tree species in the region. Species turnover was high (80%) between 6‐ and 12‐month‐old seedlings, though the /russula‐lactarius, /clavulina, and /tomentella‐thelephora lineages were species‐rich on seedlings at all ages. The number of ectomycorrhizal morphotypes per seedling increased with age, but extent of fungal colonization did not. Seedling ectomycorrhizal fungi were shared with sympatric conspecific adults (55%) and, to a lesser extent, regional heterospecific adults (27%), but numerous species were previously unrecorded for Guyana. Over their developmentD. corymbosaseedlings did not rely strictly on adult trees for their mycobionts but appeared to foster unique assemblages of ectomycorrhizal fungi.

     
    more » « less
  5. Abstract

    Large terrestrial herbivorous mammals (LTH‐mammals) influence plant community structure by affecting seedling establishment in mature tropical forests. Many of these LTH‐mammals frequent secondary forests, but their effects on seedling establishment in them are understudied, hindering our understanding of how LTH‐mammals influence forest regeneration in human‐modified landscapes.

    We tested the hypothesis that the strength of LTH‐mammals' effects on seedling establishment depends on landscape protection, forest successional stage and plant species' traits using a manipulative field experiment in six 1‐ha sites with varying successional age and landscape protection. In each site, we established 40 seedling plot‐pairs, with one plot excluding LTH‐mammals and one not, and monitored seedlings of 116 woody species for 26 months.

    We found significant effects of LTH‐mammal exclusion on seedling survival contingent upon the protection of forests at the landscape level and forest stage. After 26 months, survival differences between LTH‐mammal exclusion and non‐exclusion treatments were greater in protected than unprotected landscapes. Additionally, plant species' traits were related to the LTH‐mammals' differential effects, as LTH‐mammals reduced the survival of seedlings of larger‐seeded species the most. Overall, LTH‐mammals' effects translated into significant shifts in community composition as seedling communities inside and outside the exclosures diverged. Moreover, lower density and higher species diversity were found as early as 12 and 18 months outside than inside exclosures.

    Synthesis and applications.Insight into the interactions between LTH‐mammals and seedling communities in forest regeneration can be instrumental in planning effective restoration efforts. We highlight the importance of landscape protection in seedling survival and the role of LTH‐mammals in promoting seedling diversity in mature forests but also in secondary successional forests. The findings suggest that conservation efforts and possibly trophic rewilding can be important approaches for preserving diversity and influencing the trajectory of secondary tropical forest succession. However, we also caution that an overabundance of LTH‐mammals may adversely impact the pace of forest succession due to their preference for large‐seeded species. Therefore, a comprehensive wildlife management plan is indispensable. Additionally, longer term studies on LTH‐mammals are necessary to understand the effects of temporal fluctuations that are undetected in short‐term studies.

     
    more » « less