Long-term changes in avian biomass and functional diversity within disturbed and undisturbed Amazonian rainforest.

Recent long-term studies in protected areas have revealed the loss of biodiversity, yet the ramifications for ecosystem health and resilience remain unknown. Here, we investigate how the loss of understory birds, in the lowest stratum of the forest, affects avian biomass and functional diversity in the Amazon rainforest. Across approximately 30 years in the Biological Dynamics of Forest Fragments Project, we used a historical baseline of avian communities to contrast the avian communities in today's primary forest with those in modern disturbed habitat. We found that in primary rainforest, the reduced abundance of insectivorous species led to reduced functional diversity, but no reduction of biomass, indicating that species with similar functional traits are less likely to coexist in modern primary forests. Because today's forests contain fewer functionally redundant species-those with similar traits-we argue that avian communities in modern primary Amazonian rainforests are less resilient, which may ultimately disrupt the ecosystem in dynamic and unforeseen ways.

Long-term change in the avifauna of undisturbed Amazonian rainforest: ground-foraging birds disappear and the baseline shifts.

How are rainforest birds faring in the Anthropocene? We use bird captures spanning > 35 years from 55 sites within a vast area of intact Amazonian rainforest to reveal reduced abundance of terrestrial and near-ground insectivores in the absence of deforestation, edge effects or other direct anthropogenic landscape change. Because undisturbed forest includes far fewer terrestrial and near-ground insectivores than it did historically, today's fragments and second growth are more impoverished than shown by comparisons with modern 'control' sites. Any goals for bird community recovery in Amazonian second growth should recognise that a modern bird community will inevitably differ from a baseline from > 35 years ago. Abundance patterns driven by landscape change may be the most conspicuous manifestation of human activity, but biodiversity declines in undisturbed forest represent hidden losses, possibly driven by climate change, that may be pervasive in intact Amazonian forests and other systems considered to be undisturbed.

Nature, COVID-19, disease prevention, and climate change.

When COVID-19 arrived, usual human activity around the world paused or slowed, and nature responded to the opportunity. Even in major urban areas, people remarked on the clearer, crisper, star- brightened night skies. This issue's collection of interesting papers provides insights into ways the natural world responded, but it would seem worthwhile to take a step back and frame that phenomenon in its intellectual and epidemiological context.

Persistent effects of fragmentation on tropical rainforest canopy structure after 20 yr of isolation.

Assessing the persistent impacts of fragmentation on aboveground structure of tropical forests is essential to understanding the consequences of land use change for carbon storage and other ecosystem functions. We investigated the influence of edge distance and fragment size on canopy structure, aboveground woody biomass (AGB), and AGB turnover in the Biological Dynamics of Forest Fragments Project (BDFFP) in central Amazon, Brazil, after 22+ yr of fragment isolation, by combining canopy variables collected with portable canopy profiling lidar and airborne laser scanning surveys with long-term forest inventories. Forest height decreased by 30% at edges of large fragments (>10 ha) and interiors of small fragments (<3 ha). In larger fragments, canopy height was reduced up to 40 m from edges. Leaf area density profiles differed near edges: the density of understory vegetation was higher and midstory vegetation lower, consistent with canopy reorganization via increased regeneration of pioneers following post-fragmentation mortality of large trees. However, canopy openness and leaf area index remained similar to control plots throughout fragments, while canopy spatial heterogeneity was generally lower at edges. AGB stocks and fluxes were positively related to canopy height and negatively related to spatial heterogeneity. Other forest structure variables typically used to assess the ecological impacts of fragmentation (basal area, density of individuals, and density of pioneer trees) were also related to lidar-derived canopy surface variables. Canopy reorganization through the replacement of edge-sensitive species by disturbance-tolerant ones may have mitigated the biomass loss effects due to fragmentation observed in the earlier years of BDFFP. Lidar technology offered novel insights and observational scales for analysis of the ecological impacts of fragmentation on forest structure and function, specifically aboveground biomass storage.

Global mismatches in aboveground and belowground biodiversity.

Human activities are accelerating global biodiversity change and have resulted in severely threatened ecosystem services. A large proportion of terrestrial biodiversity is harbored by soil, but soil biodiversity has been omitted from many global biodiversity assessments and conservation actions, and understanding of global patterns of soil biodiversity remains limited. In particular, the extent to which hotspots and coldspots of aboveground and soil biodiversity overlap is not clear. We examined global patterns of these overlaps by mapping indices of aboveground (mammals, birds, amphibians, vascular plants) and soil (bacteria, fungi, macrofauna) biodiversity that we created using previously published data on species richness. Areas of mismatch between aboveground and soil biodiversity covered 27% of Earth's terrestrial surface. The temperate broadleaf and mixed forests biome had the highest proportion of grid cells with high aboveground biodiversity but low soil biodiversity, whereas the boreal and tundra biomes had intermediate soil biodiversity but low aboveground biodiversity. While more data on soil biodiversity are needed, both to cover geographic gaps and to include additional taxa, our results suggest that protecting aboveground biodiversity may not sufficiently reduce threats to soil biodiversity. Given the functional importance of soil biodiversity and the role of soils in human well-being, soil biodiversity should be considered further in policy agendas and conservation actions by adapting management practices to sustain soil biodiversity and considering soil biodiversity when designing protected areas.

Disparidades Mundiales entre la Biodiversidad Sobre y Bajo el Suelo Resumen Las actividades humanas están acelerando el cambio en la biodiversidad mundial y han tenido como resultado unos servicios ambientales severamente amenazados. Una gran proporción de la biodiversidad terrestre está albergada en el suelo, pero la biodiversidad de este ha sido omitida de varias evaluaciones mundiales de biodiversidad y de las acciones de conservación, además de que el entendimiento de los patrones mundiales de la biodiversidad del suelo permanece limitado; particularmente, la extensión del traslape entre los puntos fríos y calientes de biodiversidad sobre y bajo suelo no está clara. Examinamos los patrones mundiales de estos traslapes mapeando los índices de biodiversidad sobre el suelo (mamíferos, aves, anfibios y plantas vasculares) y bajo el suelo (bacterias, hongos y macrofauna) que creamos con datos previamente publicados de la riqueza de especies. Las áreas de disparidad entre la biodiversidad sobre y bajo el suelo cubrieron el 27% de la superficie terrestre del planeta. El bioma de los bosques templados de plantas frondosas y mixtas tuvo la proporción más alta de celdas de cuadrícula con una biodiversidad alta sobre el suelo, pero baja para en el subsuelo, mientras que los biomas boreales y de la tundra tuvieron una biodiversidad intermedia bajo el suelo, pero baja para el sobre suelo. Aunque se requieren más datos sobre la biodiversidad del suelo, tanto para cubrir los vacíos geográficos como para incluir a taxones adiciones, nuestros resultados sugieren que la protección a la biodiversidad sobre el suelo puede no reducir suficientemente las amenazas para la biodiversidad del suelo. Dada la importancia funcional de la biodiversidad del suelo y el papel de los suelos en el bienestar humano, se debería considerar a la biodiversidad del suelo mucho más en las agendas políticas y en las acciones de conservación, adaptando a las prácticas de manejo para que mantengan a la biodiversidad del suelo y la consideren cuando designen áreas protegidas.

Primary forests are irreplaceable for sustaining tropical biodiversity.

Human-driven land-use changes increasingly threaten biodiversity, particularly in tropical forests where both species diversity and human pressures on natural environments are high. The rapid conversion of tropical forests for agriculture, timber production and other uses has generated vast, human-dominated landscapes with potentially dire consequences for tropical biodiversity. Today, few truly undisturbed tropical forests exist, whereas those degraded by repeated logging and fires, as well as secondary and plantation forests, are rapidly expanding. Here we provide a global assessment of the impact of disturbance and land conversion on biodiversity in tropical forests using a meta-analysis of 138 studies. We analysed 2,220 pairwise comparisons of biodiversity values in primary forests (with little or no human disturbance) and disturbed forests. We found that biodiversity values were substantially lower in degraded forests, but that this varied considerably by geographic region, taxonomic group, ecological metric and disturbance type. Even after partly accounting for confounding colonization and succession effects due to the composition of surrounding habitats, isolation and time since disturbance, we find that most forms of forest degradation have an overwhelmingly detrimental effect on tropical biodiversity. Our results clearly indicate that when it comes to maintaining tropical biodiversity, there is no substitute for primary forests.

Evolutionary heritage influences Amazon tree ecology.

Lineages tend to retain ecological characteristics of their ancestors through time. However, for some traits, selection during evolutionary history may have also played a role in determining trait values. To address the relative importance of these processes requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate the combined influences of evolutionary heritage and selection in determining trait variation. We used trait data related to the major axes of life-history variation among tropical trees (e.g. growth and mortality rates) from 577 inventory plots in closed-canopy forest, mapped onto a phylogenetic hypothesis spanning more than 300 genera including all major angiosperm clades to test for evolutionary constraints on traits. We found significant phylogenetic signal (PS) for all traits, consistent with evolutionarily related genera having more similar characteristics than expected by chance. Although there is also evidence for repeated evolution of pioneer and shade tolerant life-history strategies within independent lineages, the existence of significant PS allows clearer predictions of the links between evolutionary diversity, ecosystem function and the response of tropical forests to global change.

Variation in stem mortality rates determines patterns of above-ground biomass in Amazonian forests: implications for dynamic global vegetation models.

Understanding the processes that determine above-ground biomass (AGB) in Amazonian forests is important for predicting the sensitivity of these ecosystems to environmental change and for designing and evaluating dynamic global vegetation models (DGVMs). AGB is determined by inputs from woody productivity [woody net primary productivity (NPP)] and the rate at which carbon is lost through tree mortality. Here, we test whether two direct metrics of tree mortality (the absolute rate of woody biomass loss and the rate of stem mortality) and/or woody NPP, control variation in AGB among 167 plots in intact forest across Amazonia. We then compare these relationships and the observed variation in AGB and woody NPP with the predictions of four DGVMs. The observations show that stem mortality rates, rather than absolute rates of woody biomass loss, are the most important predictor of AGB, which is consistent with the importance of stand size structure for determining spatial variation in AGB. The relationship between stem mortality rates and AGB varies among different regions of Amazonia, indicating that variation in wood density and height/diameter relationships also influences AGB. In contrast to previous findings, we find that woody NPP is not correlated with stem mortality rates and is weakly positively correlated with AGB. Across the four models, basin-wide average AGB is similar to the mean of the observations. However, the models consistently overestimate woody NPP and poorly represent the spatial patterns of both AGB and woody NPP estimated using plot data. In marked contrast to the observations, DGVMs typically show strong positive relationships between woody NPP and AGB. Resolving these differences will require incorporating forest size structure, mechanistic models of stem mortality and variation in functional composition in DGVMs.

Long-term changes in liana abundance and forest dynamics in undisturbed Amazonian forests.

Lianas (climbing woody vines) are important structural parasites of tropical trees and may be increasing in abundance in response to global-change drivers. We assessed long-term (-14-year) changes in liana abundance and forest dynamics within 36 1-ha permanent plots spanning -600 km2 of undisturbed rainforest in central Amazonia. Within each plot, we counted each liana stem (> or = 2 cm diameter) and measured its diameter at 1.3 m height, and then used these data to estimate liana aboveground biomass. An initial liana survey was completed in 1997-1999 and then repeated in 2012, using identical methods. Liana abundance in the plots increased by an average of 1.00% +/- 0.88% per year, leading to a highly significant (t = 6.58, df = 35, P < 0.00001) increase in liana stem numbers. Liana biomass rose more slowly over time (0.32% +/- 1.37% per year) and the mean difference between the two sampling intervals was nonsignificant (t = 1.46, df = 35, P = 0.15; paired t tests). Liana size distributions shifted significantly (chi2 = 191, df = 8, P < 0.0001; Chi-square test for independence) between censuses, mainly as a result of a nearly 40% increase in the number of smaller (2-3 cm diameter) lianas, suggesting that lianas recruited rapidly during the study. We used long-term data on rainfall and forest dynamics from our study site to test hypotheses about potential drivers of change in liana communities. Lianas generally increase with rainfall seasonality, but we found no significant trends over time (1997-2012) in five rainfall parameters (total annual rainfall, dry-season rainfall, wet-season rainfall, number of very dry months, CV of monthly rainfall). However, rates of tree mortality and recruitment have increased significantly over time in our plots, and general linear mixed-effect models suggested that lianas were more abundant at sites with higher tree mortality and flatter topography. Rising concentrations of atmospheric CO2, which may stimulate liana growth, might also have promoted liana increases. Our findings clearly support the view that lianas are increasing in abundance in old-growth tropical forests, possibly in response to accelerating forest dynamics and rising CO2 concentrations. The aboveground biomass of trees was lowest in plots with abundant lianas, suggesting that lianas could reduce forest carbon storage and potentially alter forest dynamics if they continue to proliferate.

Enhancing conservation, ecosystem services, and local livelihoods through a wildlife premium mechanism.

We propose the wildlife premium mechanism as an innovation to conserve endangered large vertebrates. The performance-based payment scheme would allow stakeholders in lower-income countries to generate revenue by recovering and maintaining threatened fauna that can also serve as umbrella species (i.e., species whose protection benefits other species with which they co-occur). There are 3 possible options for applying the premium: option 1, embed premiums in a carbon payment; option 2, link premiums to a related carbon payment, but as independent and legally separate transactions; option 3, link premiums to noncarbon payments for conserving ecosystem services (PES). Each option presents advantages, such as incentive payments to improve livelihoods of rural poor who reside in or near areas harboring umbrella species, and challenges, such as the establishment of a subnational carbon credit scheme. In Kenya, Peru, and Nepal pilot premium projects are now underway or being finalized that largely follow option 1. The Kasigau (Kenya) project is the first voluntary carbon credit project to win approval from the 2 leading groups sanctioning such protocols and has already sold carbon credits totaling over $1.2 million since June 2011. A portion of the earnings is divided among community landowners and projects that support community members and has added over 350 jobs to the local economy. All 3 projects involve extensive community management because they occur on lands where locals hold the title or have a long-term lease from the government. The monitoring, reporting, and verification required to make premium payments credible to investors include transparent methods for collecting data on key indices by trained community members and verification of their reporting by a biologist. A wildlife premium readiness fund would enable expansion of pilot programs needed to test options beyond those presented here.

Morphological consequences of climate change for resident birds in intact Amazonian rainforest.

Warming from climate change is expected to reduce body size of endotherms, but studies from temperate systems have produced equivocal results. Over four decades, we collected morphometric data on a nonmigratory understory bird community within Amazonian primary rainforest that is experiencing increasingly extreme climate. All 77 species showed lower mean mass since the early 1980s­nearly half with 95% confidence. A third of species concomitantly increased wing length, driving a decrease in mass:wing ratio for 69% of species. Seasonal precipitation patterns were generally better than temperature at explaining morphological variation. Short-term climatic conditions affected all metrics, but time trends in wing and mass:wing remained robust even after controlling for annual seasonal conditions. We attribute these results to pressures to increase resource economy under warming. Both seasonal and long-term morphological shifts suggest response to climate change and highlight its pervasive consequences, even in the heart of the world's largest rainforest.

Pervasive alteration of tree communities in undisturbed Amazonian forests.

Amazonian rainforests are some of the most species-rich tree communities on earth. Here we show that, over the past two decades, forests in a central Amazonian landscape have experienced highly nonrandom changes in dynamics and composition. Our analyses are based on a network of 18 permanent plots unaffected by any detectable disturbance. Within these plots, rates of tree mortality, recruitment and growth have increased over time. Of 115 relatively abundant tree genera, 27 changed significantly in population density or basal area--a value nearly 14 times greater than that expected by chance. An independent, eight-year study in nearby forests corroborates these shifts in composition. Contrary to recent predictions, we observed no increase in pioneer trees. However, genera of faster-growing trees, including many canopy and emergent species, are increasing in dominance or density, whereas genera of slower-growing trees, including many subcanopy species, are declining. Rising atmospheric CO2 concentrations may explain these changes, although the effects of this and other large-scale environmental alterations remain uncertain. These compositional changes could have important impacts on the carbon storage, dynamics and biota of Amazonian forests.

Competition influences tree growth, but not mortality, across environmental gradients in Amazonia and tropical Africa.

Competition among trees is an important driver of community structure and dynamics in tropical forests. Neighboring trees may impact an individual tree's growth rate and probability of mortality, but large-scale geographic and environmental variation in these competitive effects has yet to be evaluated across the tropical forest biome. We quantified effects of competition on tree-level basal area growth and mortality for trees ≥10-cm diameter across 151 ~1-ha plots in mature tropical forests in Amazonia and tropical Africa by developing nonlinear models that accounted for wood density, tree size, and neighborhood crowding. Using these models, we assessed how water availability (i.e., climatic water deficit) and soil fertility influenced the predicted plot-level strength of competition (i.e., the extent to which growth is reduced, or mortality is increased, by competition across all individual trees). On both continents, tree basal area growth decreased with wood density and increased with tree size. Growth decreased with neighborhood crowding, which suggests that competition is important. Tree mortality decreased with wood density and generally increased with tree size, but was apparently unaffected by neighborhood crowding. Across plots, variation in the plot-level strength of competition was most strongly related to plot basal area (i.e., the sum of the basal area of all trees in a plot), with greater reductions in growth occurring in forests with high basal area, but in Amazonia, the strength of competition also varied with plot-level wood density. In Amazonia, the strength of competition increased with water availability because of the greater basal area of wetter forests, but was only weakly related to soil fertility. In Africa, competition was weakly related to soil fertility and invariant across the shorter water availability gradient. Overall, our results suggest that competition influences the structure and dynamics of tropical forests primarily through effects on individual tree growth rather than mortality and that the strength of competition largely depends on environment-mediated variation in basal area.

The uncertain future of protected lands and waters.

Protected areas are intended to safeguard biodiversity in perpetuity, yet evidence suggests that widespread legal changes undermine protected area durability and efficacy. We documented these legal changes-protected area downgrading, downsizing, and degazettement (PADDD) events-in the United States and Amazonian countries and compiled available data globally. Governments of the United States and Amazonian countries enacted 269 and 440 PADDD events, respectively. Between 1892 and 2018, 73 countries enacted 3749 PADDD events, removing 519,857 square kilometers from protection and tempering regulations in an additional 1,659,972 square kilometers; 78% of events were enacted since 2000. Most PADDD events (62%) are associated with industrial-scale resource extraction and development, suggesting that PADDD may compromise biodiversity conservation objectives. Strategic policy responses are needed to address PADDD and sustain effective protected areas.

Evolutionary diversity is associated with wood productivity in Amazonian forests.

Higher levels of taxonomic and evolutionary diversity are expected to maximize ecosystem function, yet their relative importance in driving variation in ecosystem function at large scales in diverse forests is unknown. Using 90 inventory plots across intact, lowland, terra firme, Amazonian forests and a new phylogeny including 526 angiosperm genera, we investigated the association between taxonomic and evolutionary metrics of diversity and two key measures of ecosystem function: aboveground wood productivity and biomass storage. While taxonomic and phylogenetic diversity were not important predictors of variation in biomass, both emerged as independent predictors of wood productivity. Amazon forests that contain greater evolutionary diversity and a higher proportion of rare species have higher productivity. While climatic and edaphic variables are together the strongest predictors of productivity, our results show that the evolutionary diversity of tree species in diverse forest stands also influences productivity. As our models accounted for wood density and tree size, they also suggest that additional, unstudied, evolutionarily correlated traits have significant effects on ecosystem function in tropical forests. Overall, our pan-Amazonian analysis shows that greater phylogenetic diversity translates into higher levels of ecosystem function: tropical forest communities with more distantly related taxa have greater wood productivity.