We assessed whether diversity in plant hydraulic traits can explain the observed diversity in plant responses to water stress in seasonally dry tropical forests (SDTFs).
The Ecosystem Demography ...model 2 (ED2) was updated with a trait-driven mechanistic plant hydraulic module, as well as novel drought-phenology and plant water stress schemes. Four plant functional types were parameterized on the basis of meta-analysis of plant hydraulic traits. Simulations from both the original and the updated ED2 were evaluated against 5 yr of field data from a Costa Rican SDTF site and remote-sensing data over Central America.
The updated model generated realistic plant hydraulic dynamics, such as leaf water potential and stem sap flow. Compared with the original ED2, predictions from our novel traitdriven model matched better with observed growth, phenology and their variations among functional groups. Most notably, the original ED2 produced unrealistically small leaf area index (LAI) and underestimated cumulative leaf litter. Both of these biases were corrected by the updated model. The updated model was also better able to simulate spatial patterns of LAI dynamics in Central America.
Plant hydraulic traits are intercorrelated in SDTFs. Mechanistic incorporation of plant hydraulic traits is necessary for the simulation of spatiotemporal patterns of vegetation dynamics in SDTFs in vegetation models.
The distribution of tropical forest biomass across the landscape is poorly understood, particularly in increasingly common secondary tropical forests. We studied the landscape-scale distribution of ...edaphic properties, plant community characteristics, and aboveground biomass (AGB) in secondary tropical dry forests in northwest Costa Rica. We used structural equation modeling to examine conceptual models of relationships among these factors, with data from 84 0.1 ha plots. Stand age and soils explained 33%â60% of the variation in community-weighted mean values of foliar traits including specific leaf area, foliar nitrogen, phosphorus, and δ¹³C. Aboveground biomass ranged from 1.7 to 409 Mg·haâ»Â¹ among plots between 5 and >100 years old. Stand age alone explained 46% of the variation in AGB among plots, while a model including age, soil pH, traits, and forest type explained 58%. Stand age was the most important variable explaining the distribution of AGB and community characteristics in secondary forests. We speculate that plot size, landscape heterogeneity, disturbance history, and stand dynamics contribute to the unexplained variation in AGB across the landscape.
► Biomass in seasonally dry tropical forest is globally significant but poorly characterized. ► We combined data from 44 studies of aboveground biomass in seasonally dry tropical forest. ► Mean ...annual precipitation explains 55% of variation in mature forest biomass. ► Precipitation affects the maximum biomass attained by recovering secondary forests.
Seasonally dry tropical forests (SDTFs) are globally extensive but understudied, especially from a biogeochemical perspective. Historically, much of the land covered by SDTF has been cleared for agriculture or other land uses, but forests are now regenerating in some areas. Quantifying biomass stocks in mature and regenerating SDTF is important for constructing global carbon budgets and for designing local policy and management tools designed to sequester and store carbon. We reviewed 44 published and unpublished studies that estimate aboveground biomass in SDTF around the world and examined patterns of biomass across successional stages and the climatic range in which SDTF occur. Aboveground biomass in mature SDTF ranged from 39 to 334Mgha−1. A single climatic variable, mean annual precipitation (MAP), explained over 50% of the variation in aboveground biomass. Regenerating SDTFs at the wetter end of the precipitation spectrum (1500–2000mm MAP) attained a greater biomass but did not appear to reach maximum biomass faster than SDTFs at drier sites. We used spatially explicit biomass, climate, and biome range data along with the derived precipitation-biomass relationship to estimate current and potential global SDTF biomass C stocks of 8.7 and 22.1Pg respectively.
Plants on infertile soils exhibit physiological and morphological traits that support conservative internal nutrient cycling. However, potential trade-offs among use efficiencies for N, P, and ...cations are not well explored in species-rich habitats where multiple elements may limit plant production. We examined uptake efficiency and use efficiency of N, P, K, Ca, Mg, Al, and Na in plots of regenerating tropical dry forests spanning a gradient of soil fertility. Our aim was to determine whether plant responses to multiple elements are correlated, or whether there are trade-offs among exploitation strategies across stands varying in community composition, soil quality, and successional stage. For all elements, both uptake efficiency and use efficiency decreased as availability of the corresponding element increased. Plant responses to N, Na, and Al were uncoupled from uptake and use efficiencies for P and essential base cations, which were tightly correlated. N and P use efficiencies were associated with shifts in plant species composition along the soil fertility gradient, and there was also a trend towards increasing N use efficiency with stand age. N uptake efficiency was positively correlated with the abundance of tree species that associate with ectomycorrhizal fungi. Taken together, our results suggest that successional processes and local species composition interact to regulate plant responses to availability of multiple resources. Successional tropical dry forests appear to employ different strategies to maximize response to N vs. P and K.
Much of the dry tropical forest biome has been converted to agricultural land uses over the past several centuries. However, in conservation areas such as those in the Guanacaste and Tempisque ...regions of Costa Rica, tropical dry forests are regenerating due to management practices including fire suppression. To better understand the patterns of secondary succession occurring in Costa Rican tropical dry forest, we established 60 20
×
50
m plots in mature and regenerating forests in the Sector Santa Rosa (formerly known as Parque Nacional Santa Rosa) and Palo Verde National Park. Plots were stratified into three plant communities: tropical dry oak forest (
Quercus oleoides) (SROAK), Santa Rosa tropical dry forest (SRTDF), and Palo Verde tropical dry forest (PVTDF). In these plots we measured and identified and all individuals >10
cm DBH, measured but did not identify stems <10
cm but taller than 1.3
m, counted woody seedlings (<1.3
m height) and analyzed soil chemical and physical properties.
Soil properties clearly differentiated vegetation communities and defined a gradient from rocky, siltier soils with low nutrient availability (SROAK soils) to clayey, nutrient-rich soils (PVTDF). Soils in the Santa Rosa dry forest had intermediate soil properties compared to the other two plant communities and had the highest tree species richness. Successional dynamics as assessed from plots of different age showed that the patterns of change in indices of stand structure, species richness and tree community composition varied with forest type (and hence soil properties). Forest structure (densities of stems in different size classes) recovered to levels found in mature forest within 4–5 decades in SRTDF and PVTDF, but increased with stand age in the oak forest. In all plots, we identified 135 species from 45 families. Simple and partial Mantel tests showed that across all plots, both stand age and soil properties explain variation in species composition, but that there is also unexplained spatial variation in tree community composition after accounting for spatial co-variation with soils. Additional analyses suggested that this is due to β-diversity, i.e. changes in the regional species pool from the northern (Santa Rosa) to more southern area (Palo Verde). Species composition in young stands was dominated by wind-dispersed species in SRTDF and PVTDF, and by animal-dispersed species in the oak forest. We conclude that the management strategy of fire control promotes passive regeneration of secondary dry forest in Costa Rica. However, if a specific forest composition is desired, more active restoration strategies may be necessary.
Identifying factors controlling forest productivity is critical to understanding forest‐climate change feedbacks, modelling vegetation dynamics and carbon finance schemes. However, little research ...has focused on productivity in regenerating tropical forests which are expanding in their fraction of global area have an order of magnitude larger carbon uptake rates relative to older forest.
We examined above‐ground net primary productivity (ANPP) and its components (wood production and litterfall) over 10 years in forest plots that vary in successional age, soil characteristics and species composition using band dendrometers and litterfall traps in regenerating seasonally dry tropical forests in northwestern Costa Rica.
We show that the components of ANPP are differentially driven by age and annual rainfall and that local soil variation is important. Total ANPP was explained by a combination of age, annual rainfall and soil variation. Wood production comprised 35% of ANPP on average across sites and years, and was explained by annual rainfall but not forest age. Conversely, litterfall increased with forest age and soil fertility yet was not affected by annual rainfall. In this region, edaphic variability is highly correlated with plant community composition. Thus, variation in ecosystem processes explained by soil may also be partially explained by species composition.
These results suggest that future changes in annual rainfall can alter the secondary forest carbon sink, but this effect will be buffered by the litterfall flux which varies little among years. In determining the long‐term strength of the secondary forest carbon sink, both rainfall and forest age will be critical variables to track. We also conclude that detailed understanding of local site variation in soils and plant community may be required to accurately predict the impact of changing rainfall on forest carbon uptake.
Synthesis. We show that in seasonally dry tropical forest, annual rainfall has a positive relationship with the growth of above‐ground woody tissues of trees and that droughts lead to significant reductions in above‐ground productivity. These results provide evidence for climate change—carbon cycle feedbacks in the seasonal tropics and highlight the value of longitudinal data on forest regeneration.
We measured wood production and litterfall to estimate above‐ground net primary productivity over 10 years in forest that varied in age, soils and species. We show that the components of ANPP are differentially driven by age and annual rainfall and that soils are important. These results provide evidence for climate change—carbon cycle feedbacks in the seasonal tropics.
Functional recovery of secondary tropical forests Poorter, Lourens; Rozendaal, Danaë M A; Bongers, Frans ...
Proceedings of the National Academy of Sciences - PNAS,
12/2021, Volume:
118, Issue:
49
Journal Article
Peer reviewed
Open access
One-third of all Neotropical forests are secondary forests that regrow naturally after agricultural use through secondary succession. We need to understand better how and why succession varies across ...environmental gradients and broad geographic scales. Here, we analyze functional recovery using community data on seven plant characteristics (traits) of 1,016 forest plots from 30 chronosequence sites across the Neotropics. By analyzing communities in terms of their traits, we enhance understanding of the mechanisms of succession, assess ecosystem recovery, and use these insights to propose successful forest restoration strategies. Wet and dry forests diverged markedly for several traits that increase growth rate in wet forests but come at the expense of reduced drought tolerance, delay, or avoidance, which is important in seasonally dry forests. Dry and wet forests showed different successional pathways for several traits. In dry forests, species turnover is driven by drought tolerance traits that are important early in succession and in wet forests by shade tolerance traits that are important later in succession. In both forests, deciduous and compound-leaved trees decreased with forest age, probably because microclimatic conditions became less hot and dry. Our results suggest that climatic water availability drives functional recovery by influencing the start and trajectory of succession, resulting in a convergence of community trait values with forest age when vegetation cover builds up. Within plots, the range in functional trait values increased with age. Based on the observed successional trait changes, we indicate the consequences for carbon and nutrient cycling and propose an ecologically sound strategy to improve forest restoration success.
Many studies have examined individual environmental drivers of tropical tree species distributions, but edaphic and successional gradients have not been considered simultaneously. Furthermore, ...determining how functional traits influence species distributions along these gradients may help to elucidate mechanisms behind community assembly.
To assess the influence of environmental filtering on tropical dry forest (TDF) tree species distributions, we used forest inventory data from sites with large edaphic and successional gradients in NW Costa Rica. Our goals were to determine (1) whether edaphic or successional factors are more important determinants of the abundance of individual tree species in regenerating TDF, (2) how species‐level functional traits are related to edaphic and/or successional niche associations of tree species and (3) correlations between species‐level edaphic and successional niche associations.
The distributions of 82 focal tree species were strongly driven by both edaphic and successional gradients. Overall, 94% of species responded to soil chemistry, 89% to soil texture and 94% to stand age gradients. Some functional traits were correlated with the edaphic and successional niche associations of TDF tree species. Species that specialized on soils with high total nutrient concentrations had higher foliar nutrient concentrations (nitrogen and phosphorus) and lower leaf dry matter content (LDMC). Species with wider stand age niches had lower LDMC and wood density. There were no correlations between edaphic and successional niche optima of TDF tree species.
Our results indicate that successional and edaphic gradients concurrently drive community assembly in regenerating TDF. Moreover, our work underscores the importance of considering how the functional characteristics of TDF trees dictate species distributions across environmental gradients.
A plain language summary is available for this article.
Plain Language Summary
Sensitivity of forest mortality to drought in carbon‐dense tropical forests remains fraught with uncertainty, while extreme droughts are predicted to be more frequent and intense. Here, the potential ...of temporal autocorrelation of high‐frequency variability in Landsat Enhanced Vegetation Index (EVI), an indicator of ecosystem resilience, to predict spatial and temporal variations of forest biomass mortality is evaluated against in situ census observations for 64 site‐year combinations in Costa Rican tropical dry forests during the 2015 ENSO drought. Temporal autocorrelation, within the optimal moving window of 24 months, demonstrated robust predictive power for in situ mortality (leave‐one‐out cross‐validation R2 = 0.54), which allows for estimates of annual biomass mortality patterns at 30 m resolution. Subsequent spatial analysis showed substantial fine‐scale heterogeneity of forest mortality patterns, largely driven by drought intensity and ecosystem properties related to plant water use such as forest deciduousness and topography. Highly deciduous forest patches demonstrated much lower mortality sensitivity to drought stress than less deciduous forest patches after elevation was controlled. Our results highlight the potential of high‐resolution remote sensing to “fingerprint” forest mortality and the significant role of ecosystem heterogeneity in forest biomass resistance to drought.
Temporal autocorrelation of Landsat EVI residuals, an indicator of ecosystem resilience, demonstrated robust predictive power for in situ mortality over tropical dry forests in Costa Rica. Results highlight the potential of high‐resolution remote sensing to “fingerprint” the spatial heterogeneity of forest mortality.
Forests are experiencing simultaneous changes in climate, disturbance regimes, and management, all of which affect ecosystem function. Climate change is shifting ranges and altering forest ...productivity. Disturbance regimes are changing with the potential for novel interactions among disturbance types. In some areas, forest management practices are intensifying, whereas in other areas, lower-impact ecological methods are being used. Interactions among these changing factors are likely to alter ecosystem structure and function at regional to continental scales. A macrosystems approach is essential to assessing the broadscale impacts of these changes and quantify cross-scale interactions, emergent patterns, and feedbacks. A promising line of analysis is the assimilation of data with ecosystem models to scale processes to the macrosystem and generate projections based on alternative scenarios. Analyses of these projections can characterize the range of future variability in forest function and provide information to guide policy, industry, and science in a changing world.