Justin M. Becknell

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.

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.

Nitrogen, phosphorus, and cation use efficiency in stands of regenerating tropical dry forest

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.

Legume abundance along successional and rainfall gradients in Neotropical forests

The nutrient demands of regrowing tropical forests are partly satisfied by nitrogen-fixing legume trees, but our understanding of the abundance of those species is biased towards wet tropical regions. Here we show how the abundance of Leguminosae is affected by both recovery from disturbance and large-scale rainfall gradients through a synthesis of forest inventory plots from a network of 42 Neotropical forest chronosequences. During the first three decades of natural forest regeneration, legume basal area is twice as high in dry compared with wet secondary forests. The tremendous ecological success of legumes in recently disturbed, water-limited forests is likely to be related to both their reduced leaflet size and ability to fix N2, which together enhance legume drought tolerance and water-use efficiency. Earth system models should incorporate these large-scale successional and climatic patterns of legume dominance to provide more accurate estimates of the maximum potential for natural nitrogen fixation across tropical forests.Data from 42 chronosequence sites show a geater abundance of legumes in seasonally dry forests than in wet forests, particularly during early secondary succession, probably owing to legumes’ nitrogen-fixing ability and reduced leaflet size.

Effects of soil type and light on height growth, biomass partitioning, and nitrogen dynamics on 22 species of tropical dry forest tree seedlings: Comparisons between legumes and nonlegumes

PREMISE OF THE STUDY The seedling stage is particularly vulnerable to resource limitation, with potential consequences for community composition. We investigated how light and soil variation affected early growth, biomass partitioning, morphology, and physiology of 22 tree species common in tropical dry forest, including eight legumes. Our hypothesis was that legume seedlings are better at taking advantage of increased resource availability, which contributes to their successful regeneration in tropical dry forests. METHODS We grew seedlings in a full-factorial design under two light levels in two soil types that differed in nutrient concentrations and soil moisture. We measured height biweekly and, at final harvest, biomass partitioning, internode segments, leaf carbon, nitrogen, δ13C, and δ15N. KEY RESULTS Legumes initially grew taller and maintained that height advantage over time under all experimental conditions. Legumes also had the highest final total biomass and water-use efficiency in the high-light and high-resource soil. For nitrogen-fixing legumes, the amount of nitrogen derived from fixation was highest in the richer soil. Although seed mass tended to be larger in legumes, seed size alone did not account for all the differences between legumes and nonlegumes. Both belowground and aboveground resources were limiting to early seedling growth and function. CONCLUSIONS Legumes may have a different regeneration niche, in that they germinate rapidly and grow taller than other species immediately after germination, maximizing their performance when light and belowground resources are readily available, and potentially permitting them to take advantage of high light, nutrient, and water availability at the beginning of the wet season.

Edaphic factors, successional status, and functional traits drive habitat associations of trees in naturally regenerating tropical dry forests

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.