Potential aboveground biomass increase in Brazilian Atlantic Forest fragments with climate change.

Fragmented tropical forest landscapes preserve much of the remaining biodiversity and carbon stocks. Climate change is expected to intensify droughts and increase fire hazard and fire intensities, thereby causing habitat deterioration, and losses of biodiversity and carbon stock losses. Understanding the trajectories that these landscapes may follow under increased climate pressure is imperative for establishing strategies for conservation of biodiversity and ecosystem services. Here, we used a quantitative predictive modelling approach to project the spatial distribution of the aboveground biomass density (AGB) by the end of the 21st century across the Brazilian Atlantic Forest (AF) domain. To develop the models, we used the maximum entropy method with projected climate data to 2100, based on the Intergovernmental Panel on Climate Change Representative Concentration Pathway (RCP) 4.5 from the fifth Assessment Report. Our AGB models had a satisfactory performance (area under the curve > 0.75 and p value < .05). The models projected a significant increase of 8.5% in the total carbon stock. Overall, the projections indicated that 76.9% of the AF domain would have suitable climatic conditions for increasing biomass by 2100 considering the RCP 4.5 scenario, in the absence of deforestation. Of the existing forest fragments, 34.7% are projected to increase their AGB, while 2.6% are projected to have their AGB reduced by 2100. The regions likely to lose most AGB-up to 40% compared to the baseline-are found between latitudes 13° and 20° south. Overall, although climate change effects on AGB vary latitudinally for the 2071-2100 period under the RCP 4.5 scenario, our model indicates that AGB stocks can potentially increase across a large fraction of the AF. The patterns found here are recommended to be taken into consideration during the planning of restoration efforts, as part of climate change mitigation strategies in the AF and elsewhere in Brazil.

Deciphering the enigma of undetected species, phylogenetic, and functional diversity based on Good-Turing theory.

Estimating the species, phylogenetic, and functional diversity of a community is challenging because rare species are often undetected, even with intensive sampling. The Good-Turing frequency formula, originally developed for cryptography, estimates in an ecological context the true frequencies of rare species in a single assemblage based on an incomplete sample of individuals. Until now, this formula has never been used to estimate undetected species, phylogenetic, and functional diversity. Here, we first generalize the Good-Turing formula to incomplete sampling of two assemblages. The original formula and its two-assemblage generalization provide a novel and unified approach to notation, terminology, and estimation of undetected biological diversity. For species richness, the Good-Turing framework offers an intuitive way to derive the non-parametric estimators of the undetected species richness in a single assemblage, and of the undetected species shared between two assemblages. For phylogenetic diversity, the unified approach leads to an estimator of the undetected Faith's phylogenetic diversity (PD, the total length of undetected branches of a phylogenetic tree connecting all species), as well as a new estimator of undetected PD shared between two phylogenetic trees. For functional diversity based on species traits, the unified approach yields a new estimator of undetected Walker et al.'s functional attribute diversity (FAD, the total species-pairwise functional distance) in a single assemblage, as well as a new estimator of undetected FAD shared between two assemblages. Although some of the resulting estimators have been previously published (but derived with traditional mathematical inequalities), all taxonomic, phylogenetic, and functional diversity estimators are now derived under the same framework. All the derived estimators are theoretically lower bounds of the corresponding undetected diversities; our approach reveals the sufficient conditions under which the estimators are nearly unbiased, thus offering new insights. Simulation results are reported to numerically verify the performance of the derived estimators. We illustrate all estimators and assess their sampling uncertainty with an empirical dataset for Brazilian rain forest trees. These estimators should be widely applicable to many current problems in ecology, such as the effects of climate change on spatial and temporal beta diversity and the contribution of trait diversity to ecosystem multi-functionality.

Would protecting tropical forest fragments provide carbon and biodiversity cobenefits under REDD+?

Tropical forests store vast amounts of carbon and are the most biodiverse terrestrial habitats, yet they are being converted and degraded at alarming rates. Given global shortfalls in the budgets required to prevent carbon and biodiversity loss, we need to seek solutions that simultaneously address both issues. Of particular interest are carbon-based payments under the Reducing Emissions from Deforestation and Forest Degradation (REDD+) mechanism to also conserve biodiversity at no additional cost. One potential is for REDD+ to protect forest fragments, especially within biomes where contiguous forest cover has diminished dramatically, but we require empirical tests of the strength of any carbon and biodiversity cobenefits in such fragmented systems. Using the globally threatened Atlantic Forest landscape, we measured above-ground carbon stocks within forest fragments spanning 13 to 23 442 ha in area and with different degrees of isolation. We related these stocks to tree community structure and to the richness and abundance of endemic and IUCN Red-listed species. We found that increasing fragment size has a positive relationship with above-ground carbon stock and with abundance of IUCN Red-listed species and tree community structure. We also found negative relationships between distance from large forest block and tree community structure, endemic species richness and abundance, and IUCN Red-listed species abundance. These resulted in positive congruence between carbon stocks and Red-listed species, and the abundance and richness of endemic species, demonstrating vital cobenefits. As such, protecting forest fragments in hotspots of biodiversity, particularly larger fragments and those closest to sources, offers important carbon and biodiversity cobenefits. More generally, our results suggest that macroscale models of cobenefits under REDD+ have likely overlooked key benefits at small scales, indicating the necessity to apply models that include finer-grained assessments in fragmented landscapes rather than using averaged coarse-grained cells.

Invasive alien acacias rapidly stock carbon, but threaten biodiversity recovery in young second-growth forests.

Under the UN-Decade of Ecosystem Restoration and Bonn Challenge, second-growth forest is promoted as a global solution to climate change, degradation and associated losses of biodiversity and ecosystem services. Second growth is often invaded by alien tree species and understanding how this impacts carbon stock and biodiversity recovery is key for restoration planning. We assessed carbon stock and tree diversity recovery in second growth invaded by two Acacia species and non-invaded second growth, with associated edge effects, in the Brazilian Atlantic Forest. Carbon stock recovery in non-invaded forests was threefold lower than in invaded forests. Increasingly isolated, fragmented and deforested areas had low carbon stocks when non-invaded, whereas the opposite was true when invaded. Non-invaded forests recovered threefold to sixfold higher taxonomic, phylogenetic and functional diversity than invaded forest. Higher species turnover and lower nestedness in non-invaded than invaded forests underpinned higher abundance of threatened and endemic species in non-invaded forest. Non-invaded forests presented positive relationships between carbon and biodiversity, whereas in the invaded forests we did not detect any relationship, indicating that more carbon does not equal more biodiversity in landscapes with high vulnerability to invasive acacias. To deliver on combined climate change and biodiversity goals, restoration planning and management must consider biological invasion risk. This article is part of the theme issue 'Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration'.

Human impacts as the main driver of tropical forest carbon.

Understanding the mechanisms controlling forest carbon storage is crucial to support "nature-based" solutions for climate change mitigation. We used a dataset of 892 Atlantic Forest inventories to assess the direct and indirect effects of environmental conditions, human impacts, tree community proprieties, and sampling methods on tree above-ground carbon stocks. We showed that the widely accepted drivers of carbon stocks, such as climate, soil, topography, and forest fragmentation, have a much smaller role than the forest disturbance history and functional proprieties of the Atlantic Forest. Specifically, within-forest disturbance level was the most important driver, with effect at least 30% higher than any of the environmental conditions individually. Thus, our findings suggest that the conservation of tropical carbon stocks may be dependable on, principally, avoiding forest degradation and that conservation policies focusing only on carbon may fail to protect tropical biodiversity.

Defaunation affects carbon storage in tropical forests.

Carbon storage is widely acknowledged as one of the most valuable forest ecosystem services. Deforestation, logging, fragmentation, fire, and climate change have significant effects on tropical carbon stocks; however, an elusive and yet undetected decrease in carbon storage may be due to defaunation of large seed dispersers. Many large tropical trees with sizeable contributions to carbon stock rely on large vertebrates for seed dispersal and regeneration, however many of these frugivores are threatened by hunting, illegal trade, and habitat loss. We used a large data set on tree species composition and abundance, seed, fruit, and carbon-related traits, and plant-animal interactions to estimate the loss of carbon storage capacity of tropical forests in defaunated scenarios. By simulating the local extinction of trees that depend on large frugivores in 31 Atlantic Forest communities, we found that defaunation has the potential to significantly erode carbon storage even when only a small proportion of large-seeded trees are extirpated. Although intergovernmental policies to reduce carbon emissions and reforestation programs have been mostly focused on deforestation, our results demonstrate that defaunation, and the loss of key ecological interactions, also poses a serious risk for the maintenance of tropical forest carbon storage.

The hypothesis of sympatric speciation as the dominant generator of endemism in a global hotspot of biodiversity.

Allopatric or sympatric speciation influence the degree to which closely related species coexist in different manners, altering the patterns of phylogenetic structure and turnover among and between communities. The objective of this study was to examine whether phylogenetic community structure and turnover in the Brazilian Atlantic Forest permit conclusions about the dominant process for the formation of extant angiosperm richness of tree species. Therefore, we analyzed phylogenetic community structure (MPD, MNTD) as well as taxonomic (Jaccard similarity) and phylogenetic turnover (betaMPD, betaMNTD) among and between 49 tree communities distributed among three different habitat types. Mean annual precipitation and mean annual temperature in each survey area were estimated. Phylogenetic community structure does not differ between habitat types, although MPD reduces with mean annual temperature. Jaccard similarity decreases and betaMNTD increases with spatial distance and environmental differences between study sites. Spatial distance explains the largest portions of variance in the data, indicating dispersal limitation and the spatial aggregation of recently formed taxa, as betaMNTD is related to more recent evolutionary events. betaMPD, that is related to deep evolutionary splits, shows no spatial or environmental pattern, indicating that older clades are equally distributed across the Brazilian Atlantic Forest. While similarity pattern indicates dispersal limitations, the spatial turnover of betaMNTD is consistent with a high degree of sympatric speciation generating extant diversity and endemism in the Brazilian Atlantic Forest. More comprehensive approaches are necessary to reduce spatial sampling bias, uncertainties regarding angiosperm diversification patterns and confirm sympatric speciation as the dominant generator for the formation of extant species diversity in the Brazilian Atlantic Forest.

Relación especie-área y distribución de la abundancia de especies en una comunidad vegetal de un inselberg tropical: efecto del tamaño de los parches / Species-area relation and species abundance distribution in a plant community on a tropical inselberg: effect of patch size

Resumen Aunque los inselbergs son afloramientos rocosos icónicos con un alto valor biogeográfico, poco se conoce sobre los mecanismos responsables de la estructuración de comunidades vegetales. El objetivo de esta investigación fue evaluar cómo el tamaño de los parches de vegetación influye en la relación especie-área y distribución de la abundancia de especies de una comunidad en un inselberg del Monumento Natural "Piedra La Tortuga", región Guayana, Venezuela. Por este motivo, se establecieron tres preguntas de investigación: ¿Cuál es el efecto del tamaño de los parches sobre la riqueza de especies? ¿Qué tipo de modelo especie-área (SAR) presenta mejor ajuste en esos parches de vegetación? ¿Cómo es la distribución de las abundancias de las especies (SADs) es inducida por el tamaño de los parches? Se realizó un muestreo aleatorio estratificado en parches que oscilaron entre 0.34 y 14.8 m2, totalizando 40 unidades muestrales (226 m2). Todos los individuos encontrados en los 40 parches fueron identificados a nivel de especie. La composición florística en las diferentes muestras estuvo representada por 19 familias, 22 géneros y 24 especies, de las cuales 50 % son endémicas de inselbergs y dos están amenazadas de extinción. Se identificaron dos grupos de tamaños de parches (grandes 8-15 m2 y pequeños ≤ 7.9 m2) en relación a la abundancia y composición de especies, con diferencias significativas entre los grupos. Las curvas de acumulación de especies para cada grupo de tamaño de parche muestran una tendencia contrastante con marcadas diferencias en la riqueza observada entre los grupos de tamaños de parches. Las curvas de los modelos SADs tuvieron un ajuste significativo de la serie geométrica en las dos categorías de parches. El modelo SAR de la función potencia presentó los mejores ajustes especie-área, donde el aumento del área de los parches explicó un 82 % de la variación en el aumento del número de especies. Los resultados de este estudio demuestran por primera vez como el tamaño de los parches de vegetación de un inselberg tropical tiene una fuerte influencia sobre la riqueza, distribución de la abundancia y composición de especies. Así mismo, se determinó que el modelo geométrico SAD presentó el mejor ajuste en la comunidad en función del tamaño de los parches como un indicador de recursos, donde la abundancia de una especie puede ser equivalente a una proporción del espacio ocupado. También se presume que los cambios de tamaño de los parches, podría estar asociado con la disponibilidad de nutrientes y agua, como ha sido demostrado en otros ambientes de tierras secas. En algunos estudios se ha argumentado que la variación en la composición de especies entre los perfiles de vegetación de inselbergs tropicales está condicionada principalmente por la estructura del hábitat y el déficit hídrico. Sin embargo, no se había discutido cómo el tamaño de los parches de vegetación tiene un efecto en la riqueza. Los análisis SADs y SAR pueden proporcionar explicaciones complementarias sobre la estructuración de comunidades vegetales en inselbergs.

Abstract Although inselbergs are iconic rock outcrops with a high biogeographic value, little is known about drivers responsible for the plant community assembly. The aim of this research was to evaluate how the patch size distribution of vegetation influences the species-area relationship and species abundance distribution of a community in an inselberg of the "Piedra La Tortuga" Natural Monument of the Guayana region, Venezuela. In this context, three research questions were established: What is the effect of patch size on species richness? What species-area model (SAR) has the best fit in those vegetation patches? How is the distribution of species abundances (SADs) induced by the patch size distribution? A stratified random sampling was performed in patches ranging from 0.34 to 14.8 m2, totaling 40 sampling units (226 m2). All individuals found in the 40 patches were identified at species level. The floristic composition in the different samples was represented by 19 families, 22 genera and 24 species, of which 50 % are endemic to inselbergs and two, are threatened of extinction. Two groups of patch sizes were identified (large 8-15 m2 and small ≤ 7.9 m2) in relation to the abundance and composition of species. The species accumulation curves for each patch size group show a contrasting tendency with marked differences in the observed richness among patch size groups. The curves of the SADs models had a significant adjustment of the geometric series in the two categories of patches. The SAR model of the power function presented the best species-area adjustments, where the increase in patch area accounted for 82 % of the variation in the increase in the number of species. The results of this study demonstrate for the first time how vegetation patches of a tropical inselberg have a strong influence on richness, abundance distribution and species composition. Likewise, it was determined that the SAD geometric model presented the best fit in the community as a function of patch size as a resource indicator, where the abundance of a species can be equivalent to a proportion of the space occupied. It is also presumed that changes in patch sizes could be associated with nutrient and water availability, as has been demonstrated in other dryland environments. In some studies it has been argued that variation in species composition among vegetation profiles of tropical inselbergs is mainly conditioned by habitat structure and water deficit. However, it had not been discussed how the size of patches of vegetation has an effect on richness. SADs and SAR analyzes can provide complementary explanations on community assembly in inselbergs. Rev. Biol. Trop. 66(2): 937-951. Epub 2018 June 01.