Tree demographic strategies largely overlap across succession in Neotropical wet and dry forest communities.

Secondary tropical forests play an increasingly important role in carbon budgets and biodiversity conservation. Understanding successional trajectories is therefore imperative for guiding forest restoration and climate change mitigation efforts. Forest succession is driven by the demographic strategies-combinations of growth, mortality and recruitment rates-of the tree species in the community. However, our understanding of demographic diversity in tropical tree species stems almost exclusively from old-growth forests. Here, we assembled demographic information from repeated forest inventories along chronosequences in two wet (Costa Rica, Panama) and two dry (Mexico) Neotropical forests to assess whether the ranges of demographic strategies present in a community shift across succession. We calculated demographic rates for >500 tree species while controlling for canopy status to compare demographic diversity (i.e., the ranges of demographic strategies) in early successional (0-30 years), late successional (30-120 years) and old-growth forests using two-dimensional hypervolumes of pairs of demographic rates. Ranges of demographic strategies largely overlapped across successional stages, and early successional stages already covered the full spectrum of demographic strategies found in old-growth forests. An exception was a group of species characterized by exceptionally high mortality rates that was confined to early successional stages in the two wet forests. The range of demographic strategies did not expand with succession. Our results suggest that studies of long-term forest monitoring plots in old-growth forests, from which most of our current understanding of demographic strategies of tropical tree species is derived, are surprisingly representative of demographic diversity in general, but do not replace the need for further studies in secondary forests.

Soil resistance and recovery during neotropical forest succession.

The recovery of soil conditions is crucial for successful ecosystem restoration and, hence, for achieving the goals of the UN Decade on Ecosystem Restoration. Here, we assess how soils resist forest conversion and agricultural land use, and how soils recover during subsequent tropical forest succession on abandoned agricultural fields. Our overarching question is how soil resistance and recovery depend on local conditions such as climate, soil type and land-use history. For 300 plots in 21 sites across the Neotropics, we used a chronosequence approach in which we sampled soils from two depths in old-growth forests, agricultural fields (i.e. crop fields and pastures), and secondary forests that differ in age (1-95 years) since abandonment. We measured six soil properties using a standardized sampling design and laboratory analyses. Soil resistance strongly depended on local conditions. Croplands and sites on high-activity clay (i.e. high fertility) show strong increases in bulk density and decreases in pH, carbon (C) and nitrogen (N) during deforestation and subsequent agricultural use. Resistance is lower in such sites probably because of a sharp decline in fine root biomass in croplands in the upper soil layers, and a decline in litter input from formerly productive old-growth forest (on high-activity clays). Soil recovery also strongly depended on local conditions. During forest succession, high-activity clays and croplands decreased most strongly in bulk density and increased in C and N, possibly because of strongly compacted soils with low C and N after cropland abandonment, and because of rapid vegetation recovery in high-activity clays leading to greater fine root growth and litter input. Furthermore, sites at low precipitation decreased in pH, whereas sites at high precipitation increased in N and decreased in C : N ratio. Extractable phosphorus (P) did not recover during succession, suggesting increased P limitation as forests age. These results indicate that no single solution exists for effective soil restoration and that local site conditions should determine the restoration strategies. This article is part of the theme issue 'Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration'.

Strong floristic distinctiveness across Neotropical successional forests.

Forests that regrow naturally on abandoned fields are important for restoring biodiversity and ecosystem services, but can they also preserve the distinct regional tree floras? Using the floristic composition of 1215 early successional forests (≤20 years) in 75 human-modified landscapes across the Neotropic realm, we identified 14 distinct floristic groups, with a between-group dissimilarity of 0.97. Floristic groups were associated with location, bioregions, soil pH, temperature seasonality, and water availability. Hence, there is large continental-scale variation in the species composition of early successional forests, which is mainly associated with biogeographic and environmental factors but not with human disturbance indicators. This floristic distinctiveness is partially driven by regionally restricted species belonging to widespread genera. Early secondary forests contribute therefore to restoring and conserving the distinctiveness of bioregions across the Neotropical realm, and forest restoration initiatives should use local species to assure that these distinct floras are maintained.

Multidimensional tropical forest recovery.

Tropical forests disappear rapidly because of deforestation, yet they have the potential to regrow naturally on abandoned lands. We analyze how 12 forest attributes recover during secondary succession and how their recovery is interrelated using 77 sites across the tropics. Tropical forests are highly resilient to low-intensity land use; after 20 years, forest attributes attain 78% (33 to 100%) of their old-growth values. Recovery to 90% of old-growth values is fastest for soil (<1 decade) and plant functioning (<2.5 decades), intermediate for structure and species diversity (2.5 to 6 decades), and slowest for biomass and species composition (>12 decades). Network analysis shows three independent clusters of attribute recovery, related to structure, species diversity, and species composition. Secondary forests should be embraced as a low-cost, natural solution for ecosystem restoration, climate change mitigation, and biodiversity conservation.

Functional recovery of secondary tropical forests.

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.

Forest cover mediates large and medium-sized mammal occurrence in a critical link of the Mesoamerican Biological Corridor.

Connectivity of natural areas through biological corridors is essential for ecosystem resilience and biodiversity conservation. However, robust assessments of biodiversity in corridor areas are often hindered by logistical constraints and the statistical challenges of modeling data from multiple species. Herein, we used a hierarchical community occupancy model in a Bayesian framework to evaluate the status of medium and large-sized mammals in a critical link of the Mesoamerican Biological Corridor (MBC) in Costa Rica. We used camera traps deployed from 2013-2017 to detect 18 medium (1-15 kg) and 6 large (>15 kg) mammal species in a portion of two Jaguar Conservation Units (JCUs) and the Corridor linking them. Camera traps operated for 16,904 trap nights across 209 stations, covering an area of 880 km2. Forest cover was the most important driver of medium and large-sized mammal habitat use, with forest specialists such as jaguars (Panthera onca) and pumas (Puma concolor) strongly associated with high forest cover, while habitat generalists such as coyotes (Canis latrans) and raccoons (Procyon lotor) were associated with low forest cover. Medium and large-sized mammal species richness was lower in the Corridor area ([Formula: see text] = 9.78±1.84) than in the portions evaluated of the two JCUs ([Formula: see text] = 11.50±1.52). Puma and jaguar habitat use probabilities were strongly correlated with large prey species richness (jaguar, r = 0.59, p<0.001; puma, r = 0.72, p<0.001), and correlated to a lesser extent with medium prey species richness (jaguar, r = 0.36, p = 0.003; puma, r = 0.23, p = 0.064). Low estimated jaguar habitat use probability in one JCU (Central Volcanic Cordillera: [Formula: see text] = 0.15±0.11) suggests that this is not the jaguar stronghold previously assumed. In addition, the western half of the Corridor has low richness of large mammals, making it necessary to take urgent actions to secure habitat connectivity for mammal populations.

Drivers of tropical rainforest composition and alpha diversity patterns over a 2,520 m altitudinal gradient.

AIM: We sought to determine the relationship of forest composition and alpha diversity (the species diversity of a local assemblage) to altitude, soil, and spatial factors over a 440-2,950 m a.s.l gradient. LOCATION: Altitudinal gradient on the Caribbean slope of the Talamanca Cordillera, Costa Rica. TAXON: Angiosperm and gymnosperm trees, palms, and tree ferns. METHODS: We measured and identified all stems ≥10 cm dbh in 32 0.25-ha undisturbed rain forest plots over the gradient. We determined compositional patterns using Non-Metric Multidimensional Scaling (NMS) ordination, and used linear regressions to explore the relationship between four alpha diversity metrics and altitude. With variation partitioning (VARPART), we determined the compositional variation explained by altitude, soil, and spatial variables quantified using Principle Components of Neighbor matrices. RESULTS: We identified 425 species. NMS axis 1 separated a lowland zone (440-1,120 m asl) from a transitional one dominated by holarctic Oreomunnea mexicana (1,400-1,600 m asl) and Quercus-dominated forests at altitudes >2,100 m asl. The lowland zone was separated into two clusters of plots on NMS axis 2, the first in the 430-620 m asl range and the second at 1,000-1,120 masl. Regressions showed that all alpha diversity metrics were strongly negatively related to altitude (R 2 > 0.78). Overall, adjusted R 2 from VARPART was 0.43, with 0.30, 0.21, and 0.17 for altitude, soil, and space respectively. The respective adjusted R 2 of individual matrices, on controlling for the other two, was 0.06, 0.05 and 0.09 (p < 0.001). MAIN CONCLUSIONS: There are two well-defined forest compositional zones on this gradient-lowlands 430-1,120 m asl and montane forests >2,150 m asl-with a transitional zone at 1,400-1,600 m asl, where lowland tropical and montane holarctic species are found together. Montane forests are very distinct in their composition and low alpha diversity. Vegetation and soil respond to altitude, and therefore temperature, as an integrated system, a model that goes beyond niche assembly as shown by the significant effect of space in the VARPART.

Wet and dry tropical forests show opposite successional pathways in wood density but converge over time.

Tropical forests are converted at an alarming rate for agricultural use and pastureland, but also regrow naturally through secondary succession. For successful forest restoration, it is essential to understand the mechanisms of secondary succession. These mechanisms may vary across forest types, but analyses across broad spatial scales are lacking. Here, we analyse forest recovery using 1,403 plots that differ in age since agricultural abandonment from 50 sites across the Neotropics. We analyse changes in community composition using species-specific stem wood density (WD), which is a key trait for plant growth, survival and forest carbon storage. In wet forest, succession proceeds from low towards high community WD (acquisitive towards conservative trait values), in line with standard successional theory. However, in dry forest, succession proceeds from high towards low community WD (conservative towards acquisitive trait values), probably because high WD reflects drought tolerance in harsh early successional environments. Dry season intensity drives WD recovery by influencing the start and trajectory of succession, resulting in convergence of the community WD over time as vegetation cover builds up. These ecological insights can be used to improve species selection for reforestation. Reforestation species selected to establish a first protective canopy layer should, among other criteria, ideally have a similar WD to the early successional communities that dominate under the prevailing macroclimatic conditions.

Biodiversity recovery of Neotropical secondary forests.

Old-growth tropical forests harbor an immense diversity of tree species but are rapidly being cleared, while secondary forests that regrow on abandoned agricultural lands increase in extent. We assess how tree species richness and composition recover during secondary succession across gradients in environmental conditions and anthropogenic disturbance in an unprecedented multisite analysis for the Neotropics. Secondary forests recover remarkably fast in species richness but slowly in species composition. Secondary forests take a median time of five decades to recover the species richness of old-growth forest (80% recovery after 20 years) based on rarefaction analysis. Full recovery of species composition takes centuries (only 34% recovery after 20 years). A dual strategy that maintains both old-growth forests and species-rich secondary forests is therefore crucial for biodiversity conservation in human-modified tropical landscapes.

Disturbance and the elevation ranges of woody plant species in the mountains of Costa Rica.

AIM: To understand how disturbance-here defined as a transient reduction in competition-can shape plant distributions along elevation gradients. Theory suggests that disturbance may increase elevation ranges, especially at the lower range limits, through reduced competitive exclusion. Nevertheless, to date this relationship remains unclear. LOCATION: Mountains of Costa Rica. METHODS: We compared the elevation range of woody stems over 10 cm dbh ("trees") observed in plots along two transects spanning a range of elevations in secondary (regrowth) and old-growth forest (409 and 249 species, respectively). We also estimated these elevation ranges using nationwide data. In addition, we examined the influence of stem size and plot scale basal area (as a measure of competition) on species elevation range limits in the two gradients. RESULTS: In general, tree species ranges increased with elevation. Species in the secondary forest had broader elevation ranges (100-318 m broader than species in the old-growth forest; Wilcoxon: p-value <.001). Also, in the secondary transect, individuals with greater diameters had broader elevation ranges than those observed as smaller trees (137 m broader; Kruskal-Wallis: p-value = .03). The lower range limit of species occurred more frequently in plots with lower (vs. higher) basal area than expected by chance in both forest types. We also observed higher elevation upper limits in old growth, but not in secondary forests, with lower (vs. higher) basal area. MAIN CONCLUSION: Disturbance relaxes the constraints imposed by competition and extends effective elevation ranges of species, particularly those in secondary forest, to warmer and cooler climates (minimum increase equivalent to about 0.6-1.4°C). Thus, suitable disturbance may assist species persistence under climate change. We believe this is the first study indicating a consistent relation between disturbance and woody plant species distributions along elevation gradients.

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.

Comparative landscape genetics of two frugivorous bats in a biological corridor undergoing agricultural intensification.

Agricultural intensification in tropical landscapes poses a new threat to the ability of biological corridors to maintain functional connectivity for native species. We use a landscape genetics approach to evaluate impacts of expanding pineapple plantations on two widespread and abundant frugivorous bats in a biological corridor in Costa Rica. We hypothesize that the larger, more mobile Artibeus jamaicensis will be less impacted by pineapple than the smaller Carollia castanea. In 2012 and 2013, we sampled 735 bats in 26 remnant forest patches surrounded by different proportions of forest, pasture, crops and pineapple. We used 10 microsatellite loci for A. jamaicensis and 16 microsatellite loci for C. castanea to estimate genetic diversity and gene flow. Canonical correspondence analyses indicate that land cover type surrounding patches has no impact on genetic diversity of A. jamaicensis. However, for C. castanea, both percentage forest and pineapple surrounding patches explained a significant proportion of the variation in genetic diversity. Least-cost transect analyses (LCTA) and pairwise G″st suggest that for A. jamaicensis, pineapple is more permeable to gene flow than expected, while as expected, forest is the most permeable land cover for gene flow of C. castanea. For both species, LCTA indicate that development may play a role in inhibiting gene flow. The current study answers the call for landscape genetic research focused on tropical and agricultural landscapes, highlights the value of comparative landscape genetics in biological corridor design and management and is one of the few studies of biological corridors in any ecosystem to implement a genetic approach to test corridor efficacy.

Successional dynamics in Neotropical forests are as uncertain as they are predictable.

Although forest succession has traditionally been approached as a deterministic process, successional trajectories of vegetation change vary widely, even among nearby stands with similar environmental conditions and disturbance histories. Here, we provide the first attempt, to our knowledge, to quantify predictability and uncertainty during succession based on the most extensive long-term datasets ever assembled for Neotropical forests. We develop a novel approach that integrates deterministic and stochastic components into different candidate models describing the dynamical interactions among three widely used and interrelated forest attributes--stem density, basal area, and species density. Within each of the seven study sites, successional trajectories were highly idiosyncratic, even when controlling for prior land use, environment, and initial conditions in these attributes. Plot factors were far more important than stand age in explaining successional trajectories. For each site, the best-fit model was able to capture the complete set of time series in certain attributes only when both the deterministic and stochastic components were set to similar magnitudes. Surprisingly, predictability of stem density, basal area, and species density did not show consistent trends across attributes, study sites, or land use history, and was independent of plot size and time series length. The model developed here represents the best approach, to date, for characterizing autogenic successional dynamics and demonstrates the low predictability of successional trajectories. These high levels of uncertainty suggest that the impacts of allogenic factors on rates of change during tropical forest succession are far more pervasive than previously thought, challenging the way ecologists view and investigate forest regeneration.

Concepts and a methodology for evaluating environmental services from trees of small farms in Chiapas, México.

We propose a methodology to estimate the environmental service (ES) value of small farms in Chiapas, Mexico, involving trained farmers-promoters in field sampling. We considered the ways in which the landscape's principal organisms, the trees, contribute to ES. We proposed a species functional value (FV) index based on their functional traits and key ecological characteristics, and estimated each site's ES value using FV weighted by the dimensions and abundance of individuals in different land uses (LU). Tree contribution to carbon storage (C) was defined using species wood density and biodiversity conservation value (BD) using food and habitat provision for wildlife and species existence (non-use) value (EX). Many species and individuals had high C, as wind-dispersed species with dense wood were common, but low BD prevailed, with high BD species common only in riparian forests. Few species and fewer individuals had high EX conferred by dense wood, large size, harvesting pressure and animal dispersal, among others. High variance in value within LU types, suggested that LU is a poor estimator of ES value, and that the measurement of species FV and tree dimensions is essential. This tool accurately reflects the ecological values of farm tree cover, allowing negotiation of compensation for environmental services. This methodology could be implemented combining open-access regional traits databases and field sampling by local people, and can easily be adapted for the measurement of other ES, and to other ecological and cultural contexts.

[Tropical forest restoration in Costa Rica: the effect of several strategies on litter production, accumulation and decomposition]. / Restauración ecológica de bosques tropicales en Costa Rica: efecto de varios modelos en la producción, acumulación y descomposición de hojarasca.

Tropical forest restoration strategies have the potential to accelerate the recovery of the nutrient cycles in degraded lands. Litter production and its decomposition represent the main transfer of organic material and nutrients into the soil substrate. We evaluated litter production, accumulation on the forest floor, and its decomposition under three restoration strategies: plantation (entire area planted with trees), island (trees planted in patches of three different sizes) and control (natural regeneration) plots. We also compared restoration strategies to young secondary forest (7-9 yr). Restoration treatments were established in 50 x 50m plots in June 2004 at six sites in Southern Costa Rica. Planted tree species included two native timber species (Terminalia amazonia and Vochysia guatemalensis) interplanted with two N fixers (Erythrina poeppigiana and Inga edulis). Litter was collected every 15 days between September 2008 and August 2009 in 12 0.25m2 litter traps distributed within each plot; litter that accumulated on the soil surface was collected at four locations (0.25m2 quadrats) within each plot in February and May 2009. Total litter production in plantation (6.3Mg/ha) and secondary forest (7.3Mg/ha) did not differ, but were greater than in islands (3.5Mg/ha) and control (1.4 Mg/ha). Plantation had greatest accumulation of litter on the soil surface (10.6 Mg/ha) as compared to the other treatments (SF = 7.2; I = 6.7; C = 4.9). Secondary forest was the only treatment with a greater annual production of litter than litter accumulation on the soil surface. Carbon storage in litter was similar between plantation and secondary forest, and significantly greater than the other treatments. No differences were found for carbon concentration and storage in the soil among treatments. There was also high variability in the production and accumulation of litter and carbon among sites. Active restoration treatments accelerated the production of litter and carbon storage in comparison to areas under natural recovery. However, the nutrient cycle has not necessarily been restored under these conditions, as high litter accumulation on the soil surface indicates a low decomposition rate, which slows nutrient return to the soil.

Restauración ecológica de bosques tropicales en Costa Rica: efecto de varios modelos en la producción, acumulación y descomposición de hojarasca

Tropical forest restoration in Costa Rica: the effect of several strategies on litter production, accumulation and decomposition. Tropical forest restoration strategies have the potential to accelerate the recovery of the nutrient cycles in degraded lands. Litter production and its decomposition represent the main transfer of organic material and nutrients into the soil substrate. We evaluated litter production, accumulation on the forest floor, and its decomposition under three restoration strategies: plantation (entire area planted with trees), island (trees planted in patches of three different sizes) and control (natural regeneration) plots. We also compared restoration strategies to young secondary forest (7-9yr). Restoration treatments were established in 50x50m plots in June 2004 at six sites in Southern Costa Rica. Planted tree species included two native timber species (Terminalia amazonia and Vochysia guatemalensis) interplanted with two N fixers (Erythrina poeppigiana and Inga edulis). Litter was collected every 15 days between September 2008 and August 2009 in 12 0.25m² litter traps distributed within each plot; litter that accumulated on the soil surface was collected at four locations (0.25m² quadrats) within each plot in February and May 2009. Total litter production in plantation (6.3Mg/ha) and secondary forest (7.3Mg/ha) did not differ, but were greater than in islands (3.5Mg/ha) and control (1.4Mg/ha). Plantation had greatest accumulation of litter on the soil surface (10.6Mg/ha) as compared to the other treatments (SF=7.2; I=6.7; C=4.9). Secondary forest was the only treatment with a greater annual production of litter than litter accumulation on the soil surface. Carbon storage in litter was similar between plantation and secondary forest, and significantly greater than the other treatments. No differences were found for carbon concentration and storage in the soil among treatments. There was also high variability in the production and accumulation of litter and carbon among sites. Active restoration treatments accelerated the production of litter and carbon storage in comparison to areas under natural recovery. However, the nutrient cycle has not necessarily been restored under these conditions, as high litter accumulation on the soil surface indicates a low decomposition rate, which slows nutrient return to the soil. Rev. Biol. Trop. 59 (3): 1323-1336. Epub 2011 September 01.

Estrategias de restauración tienen el potencial de acelerar el restablecimiento del ciclo de nutrientes en áreas degradadas. En este estudio, se evaluó la producción de hojarasca, su acumulación y descomposición bajo tres tratamientos: plantación (toda la superficie plantada); islas (árboles sembrados en parches de tres tamaños) y testigo (regeneración natural). También se compararon bosques secundarios jóvenes (7-9 años). Los tratamientos fueron establecidos en parcelas de 50x50m en junio 2004 en seis sitios en el sur de Costa Rica. Las especies introducidas fueron dos maderables (Terminalia amazonia y Vochysia guatemalensis) intercaladas con dos fijadoras de nitrógeno (Erythrina poeppigiana e Inga edulis). La producción total de hojarasca en la plantación y bosque secundario no difirió significativamente, fue mayor que en las islas y el testigo. La plantación presentó mayor acumulación de hojarasca sobre el suelo. Los resultados indican una gran variabilidad entre los sitios. Ambas estrategias de restauración activas aceleraron la producción y acumulación de hojarasca en comparación con la regeneración. No obstante, eso no implica la restauración del ciclo de nutrientes. La elevada acumulación de hojarasca sobre el suelo indica baja tasa de descomposición y de retorno de nutrientes al suelo.

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.

Linking functional diversity and social actor strategies in a framework for interdisciplinary analysis of nature's benefits to society.

The crucial role of biodiversity in the links between ecosystems and societies has been repeatedly highlighted both as source of wellbeing and as a target of human actions, but not all aspects of biodiversity are equally important to different ecosystem services. Similarly, different social actors have different perceptions of and access to ecosystem services, and therefore, they have different wants and capacities to select directly or indirectly for particular biodiversity and ecosystem characteristics. Their choices feed back onto the ecosystem services provided to all parties involved and in turn, affect future decisions. Despite this recognition, the research communities addressing biodiversity, ecosystem services, and human outcomes have yet to develop frameworks that adequately treat the multiple dimensions and interactions in the relationship. Here, we present an interdisciplinary framework for the analysis of relationships between functional diversity, ecosystem services, and human actions that is applicable to specific social environmental systems at local scales. We connect the mechanistic understanding of the ecological role of diversity with its social relevance: ecosystem services. The framework permits connections between functional diversity components and priorities of social actors using land use decisions and ecosystem services as the main links between these ecological and social components. We propose a matrix-based method that provides a transparent and flexible platform for quantifying and integrating social and ecological information and negotiating potentially conflicting land uses among multiple social actors. We illustrate the applicability of our framework by way of land use examples from temperate to subtropical South America, an area of rapid social and ecological change.

Pollen dispersal and genetic structure of the tropical tree Dipteryx panamensis in a fragmented Costa Rican landscape.

In the face of widespread deforestation, the conservation of rainforest trees relies increasingly on their ability to maintain reproductive processes in fragmented landscapes. Here, we analysed nine microsatellite loci for 218 adults and 325 progeny of the tree Dipteryx panamensis in Costa Rica. Pollen dispersal distances, genetic diversity, genetic structure and spatial autocorrelation were determined for populations in four habitats: continuous forest, forest fragments, pastures adjacent to fragments and isolated pastures. We predicted longer but less frequent pollen movements among increasingly isolated trees. This pattern would lead to lower outcrossing rates for pasture trees, as well as lower genetic diversity and increased structure and spatial autocorrelation among their progeny. Results generally followed these expectations, with the shortest pollen dispersal among continuous forest trees (240 m), moderate distances for fragment (343 m) and adjacent pasture (317 m) populations, and distances of up to 2.3 km in isolated pastures (mean: 557 m). Variance around pollen dispersal estimates also increased with fragmentation, suggesting altered pollination conditions. Outcrossing rates were lower for pasture trees and we found greater spatial autocorrelation and genetic structure among their progeny, as well as a trend towards lower heterozygosity. Paternal reproductive dominance, the pollen contributions from individual fathers, did not vary among habitats, but we did document asymmetric pollen flow between pasture and adjacent fragment populations. We conclude that long-distance pollen dispersal helps maintain gene flow for D. panamensis in this fragmented landscape, but pasture and isolated pasture populations are still at risk of long-term genetic erosion.