Felipe Sinca

Amazonian functional diversity from forest canopy chemical assembly

Significance Canopy trees are keystone organisms that create habitat for an enormous array of flora and fauna and dominate carbon storage in tropical forests. Determining the functional diversity of tree canopies is, therefore, critical to understanding how tropical forests are assembled and predicting ecosystem responses to environmental change. Across the megadiverse Andes-to-Amazon corridor of Peru, we discovered a large-scale nested pattern of canopy chemical assembly among thousands of trees. This nested geographic and phylogenetic pattern within and among forest communities provides a different perspective on current and future alterations to the functioning of western Amazonian forests resulting from land use and climate change. Patterns of tropical forest functional diversity express processes of ecological assembly at multiple geographic scales and aid in predicting ecological responses to environmental change. Tree canopy chemistry underpins forest functional diversity, but the interactive role of phylogeny and environment in determining the chemical traits of tropical trees is poorly known. Collecting and analyzing foliage in 2,420 canopy tree species across 19 forests in the western Amazon, we discovered (i) systematic, community-scale shifts in average canopy chemical traits along gradients of elevation and soil fertility; (ii) strong phylogenetic partitioning of structural and defense chemicals within communities independent of variation in environmental conditions; and (iii) strong environmental control on foliar phosphorus and calcium, the two rock-derived elements limiting CO2 uptake in tropical forests. These findings indicate that the chemical diversity of western Amazonian forests occurs in a regionally nested mosaic driven by long-term chemical trait adjustment of communities to large-scale environmental filters, particularly soils and climate, and is supported by phylogenetic divergence of traits essential to foliar survival under varying environmental conditions. Geographically nested patterns of forest canopy chemical traits will play a role in determining the response and functional rearrangement of western Amazonian ecosystems to changing land use and climate.

Taxonomy and remote sensing of leaf mass per area (LMA) in humid tropical forests

Leaf mass per area (LMA) is a trait of central importance to plant physiology and ecosystem function, but LMA patterns in the upper canopies of humid tropical forests have proved elusive due to tall species and high diversity. We collected top-of-canopy leaf samples from 2873 individuals in 57 sites spread across the Neotropics, Australasia, and Caribbean and Pacific Islands to quantify environmental and taxonomic drivers of LMA variation, and to advance remote-sensing measures of LMA. We uncovered strong taxonomic organization of LMA, with species accounting for 70% of the global variance and up to 62% of the variation within a forest stand. Climate, growth habit, and site conditions are secondary contributors (1-23%) to the observed LMA patterns. Intraspecific variation in LMA averages 16%, which is a fraction of the variation observed between species. We then used spectroscopic remote sensing (400-2500 nm) to estimate LMA with an absolute uncertainty of 14-15 g/m2 (r2 = 0.85), or approximately 10% of the global mean. With radiative transfer modeling, we demonstrated the scalability of spectroscopic remote sensing of LMA to the canopy level. Our study indicates that remotely sensed patterns of LMA will be driven by taxonomic variation against a backdrop of environmental controls expressed at site and regional levels.

Targeted carbon conservation at national scales with high-resolution monitoring

Significance Land use is a principal driver of carbon emissions, either directly through land change processes such as deforestation or indirectly via transportation and industries supporting natural resource use. To minimize the effects of land use on the climate system, natural ecosystems are needed to offset gross emissions through carbon sequestration. Managing this critically important service must be achieved tactically if it is to be cost-effective. We have developed a high-resolution carbon mapping approach that can identify biogeographically explicit targets for carbon storage enhancement among all landholders within a country. Applying our approach to Perú reveals carbon threats and protections, as well as major opportunities for using ecosystems to sequester carbon. Our approach is scalable to any tropical forest country. Terrestrial carbon conservation can provide critical environmental, social, and climate benefits. Yet, the geographically complex mosaic of threats to, and opportunities for, conserving carbon in landscapes remain largely unresolved at national scales. Using a new high-resolution carbon mapping approach applied to Perú, a megadiverse country undergoing rapid land use change, we found that at least 0.8 Pg of aboveground carbon stocks are at imminent risk of emission from land use activities. Map-based information on the natural controls over carbon density, as well as current ecosystem threats and protections, revealed three biogeographically explicit strategies that fully offset forthcoming land-use emissions. High-resolution carbon mapping affords targeted interventions to reduce greenhouse gas emissions in rapidly developing tropical nations.

Amazonian landscapes and the bias in field studies of forest structure and biomass

Significance Although tropical forests absorb more carbon dioxide as biomass than any other terrestrial ecosystem, biomass estimates disagree substantially at landscape-to-regional scales. Current biomass maps rely upon field plots for extrapolations to larger scales, yet whether field plots accurately represent landscape-scale variables has not been assessed. To our knowledge, this is the first study to compare forest structural variables and aboveground biomass derived from field plots to those derived from their host landscapes using airborne 3D remote sensing. We found that typical field plots can produce substantially biased estimates and the number of plots needed to reduce this bias is impractical, positioning airborne remote sensing as a core tool for mapping forest structure and biomass across tropical landscapes. Tropical forests convert more atmospheric carbon into biomass each year than any terrestrial ecosystem on Earth, underscoring the importance of accurate tropical forest structure and biomass maps for the understanding and management of the global carbon cycle. Ecologists have long used field inventory plots as the main tool for understanding forest structure and biomass at landscape-to-regional scales, under the implicit assumption that these plots accurately represent their surrounding landscape. However, no study has used continuous, high-spatial-resolution data to test whether field plots meet this assumption in tropical forests. Using airborne LiDAR (light detection and ranging) acquired over three regions in Peru, we assessed how representative a typical set of field plots are relative to their surrounding host landscapes. We uncovered substantial mean biases (9–98%) in forest canopy structure (height, gaps, and layers) and aboveground biomass in both lowland Amazonian and montane Andean landscapes. Moreover, simulations reveal that an impractical number of 1-ha field plots (from 10 to more than 100 per landscape) are needed to develop accurate estimates of aboveground biomass at landscape scales. These biases should temper the use of plots for extrapolations of forest dynamics to larger scales, and they demonstrate the need for a fundamental shift to high-resolution active remote sensing techniques as a primary sampling tool in tropical forest biomass studies. The potential decrease in the bias and uncertainty of remotely sensed estimates of forest structure and biomass is a vital step toward successful tropical forest conservation and climate-change mitigation policy.

Airborne laser-guided imaging spectroscopy to map forest trait diversity and guide conservation

Airborne spectroscopy for forest traits The development of conservation priorities in the tropics is often hampered by the sparseness of ground data on biological diversity and the relative crudeness of larger-scale remote sensing data. Asner et al. developed airborne instrumentation to make large-scale maps of forest functional diversity across 72 million hectares of the Peruvian Andes and Amazon basin (see the Perspective by Kapos). They generated a suite of forest canopy functional trait maps from laser-guided imaging spectroscopy and used them to define distinct forest functional classes. These were then compared with government deforestation and land allocation data, which enabled an analysis of conservation threats and opportunities across the region. Science, this issue p. 385; see also p. 347 Large-scale mapping of tropical forest trait diversity offers an approach for conservation. Functional biogeography may bridge a gap between field-based biodiversity information and satellite-based Earth system studies, thereby supporting conservation plans to protect more species and their contributions to ecosystem functioning. We used airborne laser-guided imaging spectroscopy with environmental modeling to derive large-scale, multivariate forest canopy functional trait maps of the Peruvian Andes-to-Amazon biodiversity hotspot. Seven mapped canopy traits revealed functional variation in a geospatial pattern explained by geology, topography, hydrology, and climate. Clustering of canopy traits yielded a map of forest beta functional diversity for land-use analysis. Up to 53% of each mapped, functionally distinct forest presents an opportunity for new conservation action. Mapping functional diversity advances our understanding of the biosphere to conserve more biodiversity in the face of land use and climate change.

Convergence in the temperature response of leaf respiration across biomes and plant functional types.

Significance A major concern for terrestrial biosphere models is accounting for the temperature response of leaf respiration at regional/global scales. Most biosphere models incorrectly assume that respiration increases exponentially with rising temperature, with profound effects for predicted ecosystem carbon exchange. Based on a study of 231 species in 7 biomes, we find that the rise in respiration with temperature can be generalized across biomes and plant types, with temperature sensitivity declining as leaves warm. This finding indicates universally conserved controls on the temperature sensitivity of leaf metabolism. Accounting for the temperature function markedly lowers simulated respiration rates in cold biomes, which has important consequences for estimates of carbon storage in vegetation, predicted concentrations of atmospheric carbon dioxide, and future surface temperatures. Plant respiration constitutes a massive carbon flux to the atmosphere, and a major control on the evolution of the global carbon cycle. It therefore has the potential to modulate levels of climate change due to the human burning of fossil fuels. Neither current physiological nor terrestrial biosphere models adequately describe its short-term temperature response, and even minor differences in the shape of the response curve can significantly impact estimates of ecosystem carbon release and/or storage. Given this, it is critical to establish whether there are predictable patterns in the shape of the respiration–temperature response curve, and thus in the intrinsic temperature sensitivity of respiration across the globe. Analyzing measurements in a comprehensive database for 231 species spanning 7 biomes, we demonstrate that temperature-dependent increases in leaf respiration do not follow a commonly used exponential function. Instead, we find a decelerating function as leaves warm, reflecting a declining sensitivity to higher temperatures that is remarkably uniform across all biomes and plant functional types. Such convergence in the temperature sensitivity of leaf respiration suggests that there are universally applicable controls on the temperature response of plant energy metabolism, such that a single new function can predict the temperature dependence of leaf respiration for global vegetation. This simple function enables straightforward description of plant respiration in the land-surface components of coupled earth system models. Our cross-biome analyses shows significant implications for such fluxes in cold climates, generally projecting lower values compared with previous estimates.

Functional and biological diversity of foliar spectra in tree canopies throughout the Andes to Amazon region

Spectral properties of foliage express fundamental chemical interactions of canopies with solar radiation. However, the degree to which leaf spectra track chemical traits across environmental gradients in tropical forests is unknown. We analyzed leaf reflectance and transmittance spectra in 2567 tropical canopy trees comprising 1449 species in 17 forests along a 3400-m elevation and soil fertility gradient from the Amazonian lowlands to the Andean treeline. We developed quantitative links between 21 leaf traits and 400-2500-nm spectra, and developed classifications of tree taxa based on spectral traits. Our results reveal enormous inter-specific variation in spectral and chemical traits among canopy trees of the western Amazon. Chemical traits mediating primary production were tightly linked to elevational changes in foliar spectral signatures. By contrast, defense compounds and rock-derived nutrients tracked foliar spectral variation with changing soil fertility in the lowlands. Despite the effects of abiotic filtering on mean foliar spectral properties of tree communities, the spectra were dominated by phylogeny within any given community, and spectroscopy accurately classified 85-93% of Amazonian tree species. Our findings quantify how tropical tree canopies interact with sunlight, and indicate how to measure the functional and biological diversity of forests with spectroscopy.

Leaf-level photosynthetic capacity in lowland Amazonian and high-elevation Andean tropical moist forests of Peru.

We examined whether variations in photosynthetic capacity are linked to variations in the environment and/or associated leaf traits for tropical moist forests (TMFs) in the Andes/western Amazon regions of Peru. We compared photosynthetic capacity (maximal rate of carboxylation of Rubisco (Vcmax ), and the maximum rate of electron transport (Jmax )), leaf mass, nitrogen (N) and phosphorus (P) per unit leaf area (Ma , Na and Pa , respectively), and chlorophyll from 210 species at 18 field sites along a 3300-m elevation gradient. Western blots were used to quantify the abundance of the CO2 -fixing enzyme Rubisco. Area- and N-based rates of photosynthetic capacity at 25°C were higher in upland than lowland TMFs, underpinned by greater investment of N in photosynthesis in high-elevation trees. Soil [P] and leaf Pa were key explanatory factors for models of area-based Vcmax and Jmax but did not account for variations in photosynthetic N-use efficiency. At any given Na and Pa , the fraction of N allocated to photosynthesis was higher in upland than lowland species. For a small subset of lowland TMF trees examined, a substantial fraction of Rubisco was inactive. These results highlight the importance of soil- and leaf-P in defining the photosynthetic capacity of TMFs, with variations in N allocation and Rubisco activation state further influencing photosynthetic rates and N-use efficiency of these critically important forests.

Scale dependence of canopy trait distributions along a tropical forest elevation gradient

Average responses of forest foliar traits to elevation are well understood, but far less is known about trait distributional responses to elevation at multiple ecological scales. This limits our understanding of the ecological scales at which trait variation occurs in response to environmental drivers and change. We analyzed and compared multiple canopy foliar trait distributions using field sampling and airborne imaging spectroscopy along an Andes-to-Amazon elevation gradient. Field-estimated traits were generated from three community-weighting methods, and remotely sensed estimates of traits were made at three scales defined by sampling grain size and ecological extent. Field and remote sensing approaches revealed increases in average leaf mass per unit area (LMA), water, nonstructural carbohydrates (NSCs) and polyphenols with increasing elevation. Foliar nutrients and photosynthetic pigments displayed little to no elevation trend. Sample weighting approaches had little impact on field-estimated trait responses to elevation. Plot representativeness of trait distributions at landscape scales decreased with increasing elevation. Remote sensing indicated elevation-dependent increases in trait variance and distributional skew. Multiscale invariance of LMA, leaf water and NSC mark these traits as candidates for tracking forest responses to changing climate. Trait-based ecological studies can be greatly enhanced with multiscale studies made possible by imaging spectroscopy.

Reply to Adams et al.: Empirical versus process-based approaches to modeling temperature responses of leaf respiration

Reply to Adams et al. : Empirical versus process-based approaches to modeling temperature responses of leaf respiration