Susan G. Letcher

Rates of change in tree communities of secondary Neotropical forests following major disturbances

Rates of change in tree communities following major disturbances are determined by a complex set of interactions between local site factors, landscape history and structure, regional species pools and species life histories. Our analysis focuses on vegetation change following abandonment of agricultural fields or pastures, as this is the most extensive form of major disturbance in Neotropical forests. We consider five tree community attributes: stem density, basal area, species density, species richness and species composition. We describe two case studies, in northeastern Costa Rica and Chiapas, Mexico, where both chronosequence and annual tree dynamics studies are being applied. These case studies show that the rates of change in tree communities often deviate from chronosequence trends. With respect to tree species composition, sites of different ages differ more than a single site followed over time through the same age range. Dynamic changes in basal area within stands, on the other hand, generally followed chronosequence trends. Basal area accumulation was more linked with tree growth rates than with net changes in tree density due to recruitment and mortality. Stem turnover rates were poor predictors of species turnover rates, particularly at longer time-intervals. Effects of the surrounding landscape on tree community dynamics within individual plots are poorly understood, but are likely to be important determinants of species accumulation rates and relative abundance patterns.

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.

A novel statistical method for classifying habitat generalists and specialists

We develop a novel statistical approach for classifying generalists and specialists in two distinct habitats. Using a multinomial model based on estimated species relative abundance in two habitats, our method minimizes bias due to differences in sampling intensities between two habitat types as well as bias due to insufficient sampling within each habitat. The method permits a robust statistical classification of habitat specialists and generalists, without excluding rare species a priori. Based on a user-defined specialization threshold, the model classifies species into one of four groups: (1) generalist; (2) habitat A specialist; (3) habitat B specialist; and (4) too rare to classify with confidence. We illustrate our multinomial classification method using two contrasting data sets: (1) bird abundance in woodland and heath habitats in southeastern Australia and (2) tree abundance in second-growth (SG) and old-growth (OG) rain forests in the Caribbean lowlands of northeastern Costa Rica. We evaluate the multinomial model in detail for the tree data set. Our results for birds were highly concordant with a previous nonstatistical classification, but our method classified a higher fraction (57.7%) of bird species with statistical confidence. Based on a conservative specialization threshold and adjustment for multiple comparisons, 64.4% of tree species in the full sample were too rare to classify with confidence. Among the species classified, OG specialists constituted the largest class (40.6%), followed by generalist tree species (36.7%) and SG specialists (22.7%). The multinomial model was more sensitive than indicator value analysis or abundance-based phi coefficient indices in detecting habitat specialists and also detects generalists statistically. Classification of specialists and generalists based on rarefied subsamples was highly consistent with classification based on the full sample, even for sampling percentages as low as 20%. Major advantages of the new method are (1) its ability to distinguish habitat generalists (species with no significant habitat affinity) from species that are simply too rare to classify and (2) applicability to a single representative sample or a single pooled set of representative samples from each of two habitat types. The method as currently developed can be applied to no more than two habitats at a time.

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.

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.

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.

Juvenile tree growth in relation to light availability in second-growth tropical rain forests

Light is a key environmental factor limiting growth and survival of trees in the subcanopy of wet tropical forests (Davies 2001, Thomas 1996). Light availability varies both vertically and horizontally and affects tree height, crown shape and tree architecture (Bongers & Sterck 1998, Sterck & Bongers 2001, Sterck et al. 1999) in addition to growth and survival (Clark & Clark 1992, 2001). Although many studies of tree seedlings and saplings have shown that growth varies significantly with light availability in tropical wet forests (Clark et al. 1993, Iriarte & Chazdon 2005, King 1991, Kohyama 1991, Montgomery & Chazdon 2002, Oberbauer et al. 1988, 1993; Poorter & Werger 1999, Sterck et al. 1999, Welden et al. 1991), few studies have examined these relationships in size classes above 5 cm dbh (Sterck 1999). King et al. (2005) found that annual increment growth of trees in the 8-20-cm dbh size class in two Asian forests was positively dependent on an index of crown light interception, but no direct measurements of light availability were taken in this study. Due to logistical challenges, few direct measurements of light environments above tree crowns have been made in tropical forests (Sterck & Bongers 2001). To our knowledge, no measurements have been made in second-growth forests. We examined variation in growth rates and crown sizes of trees in the 5-9.9-cm dbh size class in response to variation in light availability above crowns of three species in two second-growth, 1-ha plots at La Selva Biological Station in north-eastern Costa Rica (Chazdon et al. 2005). Trees in this size class establish beneath the pioneer successional canopy and represent the future generation of canopy tree species. Gaps are infrequent in young secondary forests of this region (Montgomery & Chazdon 2001, Nicotra et al. 1999) and tree recruitment into the subcanopy and canopy occurs in the absence of gaps through continuous vertical growth through the understorey and subcanopy. The natural vegetation in the area is classified as Tropical Wet Forest (sensu Holdridge et al. 1971). Both plots are on ultisols derived from weathered volcanic deposits (Sanford et al. 1994). Stands developed on abandoned cattle pastures. In 2004, LEP was 27y old and LSUR was 19 y old; stand age was documented by aerial photographs and personal accounts. In 2004, basal area of trees > 10 cm dbh was greater in the older LEP stand than in the LSUR stand (27.2 and 22.0 m2 ha-1, respectively). We studied three shade-tolerant tree species that were present in both sites and that were 5-9.9 cm dbh in 2003, with a maximum crown height of 10 m; Ocotea leucoxylon (Sw.) Laness. (Lauraceae), Pentaclethra macroloba (Willd.) Kuntze (Fabaceae) and Virola sebifera Aubl. (Myristicaceae). These were the most abundant tree species in this size class in the two study plots. Ocotea leucoxylon is a subcanopy tree, whereas the other two species are canopy trees (Hartshorn & Hammel 1994). This study is part of a long-term project to monitor vegetation dynamics in second-growth tropical forests in north-eastern Costa Rica (Chazdon et al. 2005). In 2004, we sampled light availability for all trees present in the 5-9.9-cm dbh size class in 2003, with the exception of 10 Pentaclethra trees in LSUR that were omitted due to equipment failure. In total, 91 trees were measured (10 of 0. leucoxylon, 64 of P. macroloba and 17 of V. sebifera). We used a red:far-red (R:FR) sensor to evaluate light availability above individual tree crowns, based on the rapid assessment technique of *Current address: Ross University School of Medicine, P.O. Box 266 Roseau, Commonwealth of Dominica, West Indes.

Aboveground and belowground competition affect seedling performance and allometry in a tropical monsoon forest

Distinguishing the relative effects of above- and belowground competition can improve our understanding of the forces shaping community assembly in different ecosystems. In this study, we investigated the impacts of above- and belowground competition on seedling growth and allometry in a tropical monsoon forest (TMF) on Hainan Island, China. Four common dominant canopy tree species from three forest types (Schima superba in secondary forest, Peltophorum tonkinense and Vatica mangachapoi in lowland rainforest, and Terminalia nigrovenulosa in TMF) were planted in different treatments (understory vegetation removal and trenching to decrease above- and belowground competition, respectively). Three species had greater relative growth rates in height in the vegetation removal, but not in the trenching treatment. Vegetation removal reduced mortality rates for all species and increased biomass in three of the four. Trenching alone did not affect biomass for any species. Vegetation removal affected seedling allometry, resulting in higher leaf mass fraction in the lowland rainforest species P. tonkinense and V. mangachapoi. The secondary forest species, S. superba, did not successfully establish in plots with intact understory because of aboveground competition. The highly drought-adapted traits of T. nigrovenulosa allowed its seedlings to establish successfully in TMF, where it is a canopy dominant. Our results demonstrate that aboveground competition is more important than belowground competition in structuring seedling communities in TMF, because light availability largely limits seedling establishment in this ecosystem.

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.