Vanessa K. Boukili

Biomass resilience of Neotropical secondary forests

Land-use change occurs nowhere more rapidly than in the tropics, where the imbalance between deforestation and forest regrowth has large consequences for the global carbon cycle. However, considerable uncertainty remains about the rate of biomass recovery in secondary forests, and how these rates are influenced by climate, landscape, and prior land use. Here we analyse aboveground biomass recovery during secondary succession in 45 forest sites and about 1,500 forest plots covering the major environmental gradients in the Neotropics. The studied secondary forests are highly productive and resilient. Aboveground biomass recovery after 20 years was on average 122 megagrams per hectare (Mg ha−1), corresponding to a net carbon uptake of 3.05 Mg C ha−1 yr−1, 11 times the uptake rate of old-growth forests. Aboveground biomass stocks took a median time of 66 years to recover to 90% of old-growth values. Aboveground biomass recovery after 20 years varied 11.3-fold (from 20 to 225 Mg ha−1) across sites, and this recovery increased with water availability (higher local rainfall and lower climatic water deficit). We present a biomass recovery map of Latin America, which illustrates geographical and climatic variation in carbon sequestration potential during forest regrowth. The map will support policies to minimize forest loss in areas where biomass resilience is naturally low (such as seasonally dry forest regions) and promote forest regeneration and restoration in humid tropical lowland areas with high biomass resilience.

Carbon sequestration potential of second-growth forest regeneration in the Latin American tropics

Models reveal the high carbon mitigation potential of tropical forest regeneration. Regrowth of tropical secondary forests following complete or nearly complete removal of forest vegetation actively stores carbon in aboveground biomass, partially counterbalancing carbon emissions from deforestation, forest degradation, burning of fossil fuels, and other anthropogenic sources. We estimate the age and spatial extent of lowland second-growth forests in the Latin American tropics and model their potential aboveground carbon accumulation over four decades. Our model shows that, in 2008, second-growth forests (1 to 60 years old) covered 2.4 million km2 of land (28.1% of the total study area). Over 40 years, these lands can potentially accumulate a total aboveground carbon stock of 8.48 Pg C (petagrams of carbon) in aboveground biomass via low-cost natural regeneration or assisted regeneration, corresponding to a total CO2 sequestration of 31.09 Pg CO2. This total is equivalent to carbon emissions from fossil fuel use and industrial processes in all of Latin America and the Caribbean from 1993 to 2014. Ten countries account for 95% of this carbon storage potential, led by Brazil, Colombia, Mexico, and Venezuela. We model future land-use scenarios to guide national carbon mitigation policies. Permitting natural regeneration on 40% of lowland pastures potentially stores an additional 2.0 Pg C over 40 years. Our study provides information and maps to guide national-level forest-based carbon mitigation plans on the basis of estimated rates of natural regeneration and pasture abandonment. Coupled with avoided deforestation and sustainable forest management, natural regeneration of second-growth forests provides a low-cost mechanism that yields a high carbon sequestration potential with multiple benefits for biodiversity and ecosystem services.

Fog water and ecosystem function: heterogeneity in a California redwood forest.

Fog is thought to influence ecological function in coastal forests worldwide, yet few data are available that illuminate the mechanisms underlying this influence. In a California redwood forest we measured water and nitrogen (N) fluxes from horizontally moving fog and vertically delivered rain as well as redwood tree function. The spatial heterogeneity of water and N fluxes, water availability, tree water use, and canopy N processing varied greatly across seasons. Water and N fluxes to soil (annual average of 98% and 89%, respectively) across the whole forest occurred primarily in the rain season and was relatively even across the whole forest. In contrast, below-canopy flux of fog water and N declined exponentially from the windward edge to the forest interior. Following large fog events, soil moisture was greater at the windward edge than anywhere else in the forest. Physiological activity in redwoods reflected these differences in inputs across seasons: tree physiological responses did not vary spatially in the rain season, but in the fog season, water use was greater, yet water stress was less, in trees at the windward edge of the forest versus the interior. In both seasons, vertical passage through the forest changed the amount of water and form and concentration of N, revealing the role of the tree canopy in processing atmospheric inputs. Although total fog water inputs were comparatively small, they may have important ecosystem functions, including relief of canopy water stress and, where there is fog drip, functional coupling of above- and belowground processes.

Trait-mediated assembly processes predict successional changes in community diversity of tropical forests

Significance The assembly of ecological communities results from multiple mechanisms acting concurrently. Disentangling their relative importance represents a major challenge. Our study quantifies the relative importance of four trait-mediated mechanisms and associated changes in functional diversity over succession. We leverage the rapid community assembly in tropical successional forests and the rich information in functional trait data and spatially explicit long-term tree demographic data. Trait correlations with average species survival rates were the most important processes affecting survival and linked to a decrease in wood trait diversity during succession. Simultaneously, evidence for stabilizing niche differences associated with leaf traits was reflected in increasing leaf functional diversity. Our results suggest mechanisms by which individual performance affects successional change in community-wide diversity. Interspecific differences in relative fitness can cause local dominance by a single species. However, stabilizing interspecific niche differences can promote local diversity. Understanding these mechanisms requires that we simultaneously quantify their effects on demography and link these effects to community dynamics. Successional forests are ideal systems for testing assembly theory because they exhibit rapid community assembly. Here, we leverage functional trait and long-term demographic data to build spatially explicit models of successional community dynamics of lowland rainforests in Costa Rica. First, we ask what the effects and relative importance of four trait-mediated community assembly processes are on tree survival, a major component of fitness. We model trait correlations with relative fitness differences that are both density-independent and -dependent in addition to trait correlations with stabilizing niche differences. Second, we ask how the relative importance of these trait-mediated processes relates to successional changes in functional diversity. Tree dynamics were more strongly influenced by trait-related interspecific variation in average survival than trait-related responses to neighbors, with wood specific gravity (WSG) positively correlated with greater survival. Our findings also suggest that competition was mediated by stabilizing niche differences associated with specific leaf area (SLA) and leaf dry matter content (LDMC). These drivers of individual-level survival were reflected in successional shifts to higher SLA and LDMC diversity but lower WSG diversity. Our study makes significant advances to identifying the links between individual tree performance, species functional traits, and mechanisms of tropical forest succession.

Demographic drivers of successional changes in phylogenetic structure across life‐history stages in plant communities

To gain insight into the ecological processes driving community reassembly in disturbed ecosystems, we assessed the phylogenetic dispersion of early- and late-successional tree species occurring in lowland forests of northeastern Costa Rica. Early-successional species were more closely related than expected by chance, whereas late-successional species tended to be less closely related than expected by chance. Then, we evaluated temporal changes in the phylogenetic structure of seedling and tree assemblages in four 1-ha plots of secondary forests in this region. We found an increase in the phylogenetic evenness among tree individuals over time in all secondary tree assemblages, indicating that relatedness among tree individuals decreases as succession unfolds. This pattern was jointly promoted by recruitment and mortality processes, suggesting that increasing evenness was caused by the replacement of individuals of early-successional species from closely related lineages by late-successional species belonging to a wider diversity of lineages. Based on species occurrence, however, tree community reassembly did not show any significant phylogenetic trend over time. These results suggest that shifts in species abundance over succession have a greater impact on the phylogenetic structure of the community than the turnover of species. Seedling assemblages showed higher phylogenetic evenness than tree assemblages, suggesting that propagule colonization is an important process driving phylogenetic changes in species composition throughout succession. Overall, our findings showed that the phylogenetic structure of these successional communities varies at two temporal scales. At short timescales, decreased dominance by early-successional species over succession leads to increased evenness among tree individuals. At longer timescales, colonization processes result in increased phylogenetic evenness in seedling communities compared to tree communities, forecasting increasing phylogenetic evenness among adult individuals at late-successional stages.

Radial changes in wood specific gravity of tropical trees: inter‐ and intraspecific variation during secondary succession

Summary Variation in wood specific gravity (WSG) within and across species of tropical trees is poorly studied in relation to vegetation change during tropical forest succession. We investigated WSG of 91 species in eight long-term plots along a successional chronosequence in north-eastern Costa Rica. Radial changes in WSG were described by the rate of change (slope) along the stem radius. Significant radial changes were found in 42 of 74 species, with 37 species exhibiting increases from inner to outer wood, and five exhibiting decreases. Radial increases were commonly observed in species with inner WSG below 0·5, whereas radial decreases were observed in species with inner WSG above 0·7. Wood specific gravity weighted by cross-sectional area (wWSG) varied four-fold among species. Species classified as second-growth specialists had lower wWSG, lower inner and outer WSG, and higher slopes than old-growth specialists; successional generalists showed intermediate trait values. Among 18 species sampled in both second- and old-growth forests, four species (22%) showed significant variation in wWSG between forest types. Of 33 widely sampled species, seven species (21%) showed a significant effect of stem diameter on wWSG. Second-growth plots had lower stand-level wWSG and more pronounced radial increases than old-growth plots. Individual tree biomass for species with radial increases and low WSG was substantially underestimated when based on unweighted WSG compared with wWSG. Wood specific gravity varied with successional stage at multiple levels: within species, among successional specialist groups, and across plots of varying ages. Radial increases in WSG are common among trees in early and intermediate stages of tropical forest succession. This trait may confer growth advantages early in succession and increased resistance to physical or biotic damage during later successional stages.

Environmental gradients and the evolution of successional habitat specialization: a test case with 14 Neotropical forest sites

Successional gradients are ubiquitous in nature, yet few studies have systematically examined the evolutionary origins of taxa that specialize at different successional stages. Here we quantify successional habitat specialization in Neotropical forest trees and evaluate its evolutionary lability along a precipitation gradient. Theoretically, successional habitat specialization should be more evolutionarily conserved in wet forests than in dry forests due to more extreme microenvironmental differentiation between early and late-successional stages in wet forest. We applied a robust multinomial classification model to samples of primary and secondary forest trees from 14 Neotropical lowland forest sites spanning a precipitation gradient from 788 to 4000 mm annual rainfall, identifying species that are old-growth specialists and secondary forest specialists in each site. We constructed phylogenies for the classified taxa at each site and for the entire set of classified taxa and tested whether successional habitat specialization is phylogenetically conserved. We further investigated differences in the functional traits of species specializing in secondary vs. old-growth forest along the precipitation gradient, expecting different trait associations with secondary forest specialists in wet vs. dry forests since water availability is more limiting in dry forests and light availability more limiting in wet forests. Successional habitat specialization is non-randomly distributed in the angiosperm phylogeny, with a tendency towards phylogenetic conservatism overall and a trend towards stronger conservatism in wet forests than in dry forests. However, the specialists come from all the major branches of the angiosperm phylogeny, and very few functional traits showed any consistent relationships with successional habitat specialization in either wet or dry forests. Synthesis. The niche conservatism evident in the habitat specialization of Neotropical trees suggests a role for radiation into different successional habitats in the evolution of species-rich genera, though the diversity of functional traits that lead to success in different successional habitats complicates analyses at the community scale. Examining the distribution of particular lineages with respect to successional gradients may provide more insight into the role of successional habitat specialization in the evolution of species-rich taxa.

Fog as a source of nitrogen for redwood trees: evidence from fluxes and stable isotopes

Author(s): Templer, PH; Weathers, KC; Ewing, HA; Dawson, TE; Mambelli, S; Lindsey, AM; Webb, J; Boukili, VK; Firestone, MK | Abstract:

Opposing mechanisms affect taxonomic convergence between tree assemblages during tropical forest succession

Whether successional forests converge towards an equilibrium in species composition remains an elusive question, hampered by high idiosyncrasy in successional dynamics. Based on long-term tree monitoring in second-growth (SG) and old-growth (OG) forests in Costa Rica, we show that patterns of convergence between pairs of forest stands depend upon the relative abundance of species exhibiting distinct responses to the successional gradient. For instance, forest generalists contributed to convergence between SG and OG forests, whereas rare species and old-growth specialists were a source of divergence. Overall, opposing trends in taxonomic similarity among different subsets of species nullified each other, producing a net outcome of stasis over time. Our results offer an explanation for the limited convergence observed between pairwise communities and suggest that rare species and old-growth specialists may be prone to dispersal limitation, while the dynamics of generalists and second-growth specialists are more predictable, enhancing resilience in tropical secondary forests.