Climate change determines the sign of productivity trends in US forests.

Forests are integral to the global land carbon sink, which has sequestered ~30% of anthropogenic carbon emissions over recent decades. The persistence of this sink depends on the balance of positive drivers that increase ecosystem carbon storage-e.g., CO2 fertilization-and negative drivers that decrease it-e.g., intensifying disturbances. The net response of forest productivity to these drivers is uncertain due to the challenge of separating their effects from background disturbance-regrowth dynamics. We fit non-linear models to US forest inventory data (113,806 plot remeasurements in non-plantation forests from ~1999 to 2020) to quantify productivity trends while accounting for stand age, tree mortality, and harvest. Productivity trends were generally positive in the eastern United States, where climate change has been mild, and negative in the western United States, where climate change has been more severe. Productivity declines in the western United States cannot be explained by increased mortality or harvest; these declines likely reflect adverse climate-change impacts on tree growth. In the eastern United States, where data were available to partition biomass change into age-dependent and age-independent components, forest maturation and increasing productivity (likely due, at least in part, to CO2 fertilization) contributed roughly equally to biomass carbon sinks. Thus, adverse effects of climate change appear to overwhelm any positive drivers in the water-limited forests of the western United States, whereas forest maturation and positive responses to age-independent drivers contribute to eastern US carbon sinks. The future land carbon balance of forests will likely depend on the geographic extent of drought and heat stress.

Land conflicts from overlapping claims in Brazil's rural environmental registry.

Satellite-based land use monitoring and farm-level traceability offer opportunities for targeted zero-deforestation interventions on private lands. Brazil's Rural Environmental Registry (Cadastro Ambiental Rural, or "CAR"), a land cadaster based on self-declaration of property boundaries, was created to monitor compliance with national forest laws. It has become an important enabling measure for sustainable supply chain initiatives like the Amazon Soy Moratorium. However, CAR enrollment is increasingly used to bolster illegal land claims, putting it at the heart of land grabbing dynamics. Self-declaration of properties in the CAR offers a unique situation to study land conflicts and their impact on land use decisions on a large scale. We quantified competing land claims among 846,420 registrations in the Brazilian Legal Amazon and applied a series of generalized linear mixed-effects models. We determined that CAR overlaps are more prevalent on larger registrations, in more densely settled areas, and in areas with less secure land tenure. We tested how landholders respond to land conflicts, finding significantly more deforestation and declared legal forest reserve on lands with multiple claims. CAR overlap results in an overestimation of forest reserves by up to 9.7 million hectares when considering double-counted and deforested areas of reserves, highlighting an overlooked form of Forest Code noncompliance. While the CAR continues to be used as evidence of land tenure, we conclude that the formalization of land claims through self-declarations is inadequate to decrease conflicts. CAR overlap information provides objective evidence of land conflict that authorities can leverage with field inspection to ensure peaceful occupation before issuing land titles.

Biodiversity and productivity in eastern US forests.

Despite experimental and observational studies demonstrating that biodiversity enhances primary productivity, the best metric for predicting productivity at broad geographic extents-functional trait diversity, phylogenetic diversity, or species richness-remains unknown. Using >1.8 million tree measurements from across eastern US forests, we quantified relationships among functional trait diversity, phylogenetic diversity, species richness, and productivity. Surprisingly, functional trait and phylogenetic diversity explained little variation in productivity that could not be explained by tree species richness. This result was consistent across the entire eastern United States, within ecoprovinces, and within data subsets that controlled for biomass or stand age. Metrics of functional trait and phylogenetic diversity that were independent of species richness were negatively correlated with productivity. This last result suggests that processes that determine species sorting and packing are likely important for the relationships between productivity and biodiversity. This result also demonstrates the potential confusion that can arise when interdependencies among different diversity metrics are ignored. Our findings show the value of species richness as a predictive tool and highlight gaps in knowledge about linkages between functional diversity and ecosystem functioning.

Flower production decreases with warmer and more humid atmospheric conditions in a western Amazonian forest.

Climate models predict that everwet western Amazonian forests will face warmer and wetter atmospheric conditions, and increased cloud cover. It remains unclear how these changes will impact plant reproductive performance, such as flowering, which plays a central role in sustaining food webs and forest regeneration. Warmer and wetter nights may cause reduced flower production, via increased dark respiration rates or alteration in the reliability of flowering cue-based processes. Additionally, more persistent cloud cover should reduce the amounts of solar irradiance, which could limit flower production. We tested whether interannual variation in flower production has changed in response to fluctuations in irradiance, rainfall, temperature, and relative humidity over 18 yrs in an everwet forest in Ecuador. Analyses of 184 plant species showed that flower production declined as nighttime temperature and relative humidity increased, suggesting that warmer nights and greater atmospheric water saturation negatively impacted reproduction. Species varied in their flowering responses to climatic variables but this variation was not explained by life form or phylogeny. Our results shed light on how plant communities will respond to climatic changes in this everwet region, in which the impacts of these changes have been poorly studied compared with more seasonal Neotropical areas.

Root-associated fungal communities are influenced more by soils than by plant-host root traits in a Chinese tropical forest.

Forest fungal communities are shaped by the interactions between host tree root systems and the associated soil conditions. We investigated how the soil environment, root morphological traits, and root chemistry influence root-inhabiting fungal communities in three tropical forest sites of varying successional status in Xishuangbanna, China. For 150 trees of 66 species, we measured root morphology and tissue chemistry. Tree species identity was confirmed by sequencing rbcL, and root-associated fungal (RAF) communities were determined using high-throughput ITS2 sequencing. Using distance-based redundancy analysis and hierarchical variation partitioning, we quantified the relative importance of two soil variables (site average total phosphorus and available phosphorus), four root traits (dry matter content, tissue density, specific tip abundance, and forks), and three root tissue elemental concentrations (nitrogen, calcium, and manganese) on RAF community dissimilarity. The root and soil environment collectively explained 23% of RAF compositional variation. Soil phosphorus explained 76% of that variation. Twenty fungal taxa differentiated RAF communities among the three sites. Soil phosphorus most strongly affects RAF assemblages in this tropical forest. Variation in root calcium and manganese concentrations and root morphology among tree hosts, principally an architectural trade-off between dense, highly branched vs less-dense, herringbone-type root systems, are important secondary determinants.

The physiological acclimation and growth response of Populus trichocarpa to warming.

Plant metabolic acclimation to thermal stress remains underrepresented in current global climate models. Gaps exist in our understanding of how metabolic processes (i.e., photosynthesis, respiration) acclimate over time and how aboveground versus belowground acclimation differs. We measured the thermal acclimation of Populus trichocarpa, comparing aboveground versus belowground physiology over time. Ninety genetically identical ramets were propagated in mesocosms that separated root and microbial components. After establishment at 25°C for 6 weeks, 60 clones were warmed +4 or +8°C and monitored for 10 weeks, measuring photosynthesis (A), leaf respiration (R), soil respiration (Rs ), root plus soil respiration (Rs+r ), and root respiration (Rr ). We observed thermal acclimation in both A and R, with rates initially increasing, then declining as the thermal photosynthetic optimum (Topt ) and the temperature-sensitivity (Q10 ) of respiration adjusted to warmer conditions. Photosynthetic acclimation was constructive, based on an increase in both Topt and peak A. Belowground, Rs+r decreased linearly with warming, while Rs rates declined abruptly, then remained constant with additional warming. Plant biomass was greatest at +4°C, with 30% allocated belowground. Rates of mass-based Rr were similar among treatments; however, root nitrogen declined at +8°C leading to less mass nitrogen-based Rr in that treatment. The Q10 -temperature relationship of Rr was affected by warming, leading to differing values among treatments. Aboveground acclimation exceeded belowground acclimation, and plant nitrogen-use mediated the acclimatory response. Results suggest that moderate climate warming (+4°C) may lead to acclimation and increased plant biomass production but increases in production could be limited with severe warming (+8°C).

Temporal population variability in local forest communities has mixed effects on tree species richness across a latitudinal gradient.

Among the local processes that determine species diversity in ecological communities, fluctuation-dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness.

Proposing the solar-wind energy flux hypothesis as a driver of inter-annual variation in tropical tree reproductive effort.

PREMISE: The El Niño Southern Oscillation (ENSO) affects tropical environmental conditions, potentially altering ecosystem function as El Niño events interact with longer-term climate change. Anomalously warm equatorial Pacific Ocean temperatures affect rainfall and temperature throughout the tropics and coincide with altered leaf flush phenology and increased fruit production in wet tropical forests; however, the understanding of mechanisms underlying this pattern is limited. There is evidence that increases in tropical tree reproduction anticipate El Niño onset, motivating the continued search for a global driver of tropical angiosperm reproduction. We present the solar-wind energy flux hypothesis: that physical energy influx to the Earth's upper atmosphere and magnetosphere, generated by a positive anomaly in the solar wind preceding El Niño development, cues tropical trees to increase resource allocation to reproduction. METHODS: We test this hypothesis using 19 years of data from Luquillo, Puerto Rico, correlating them with measures of solar-wind energy. RESULTS: From 1994 to 2013, the solar-wind energy flux into Earth's magnetosphere (Ein ) was more strongly correlated with the number of species fruiting and flowering than the Niño 3.4 climate index, despite Niño 3.4 being previously identified as a driver of interannual increases in reproduction. CONCLUSIONS: Changes in the global magnetosphere and thermosphere conditions from increased solar-wind energy affect global atmospheric pressure and circulation patterns, principally by weakening the Walker circulation. We discuss the idea that these changes cue interannual increases in tropical tree reproduction and act through an unidentified mechanism that anticipates and synchronizes the reproductive output of the tropical trees with El Niño.

Understanding the recruitment response of juvenile Neotropical trees to logging intensity using functional traits.

Selective logging remains a widespread practice in tropical forests, yet the long-term effects of timber harvest on juvenile tree (i.e., sapling) recruitment across the hundreds of species occurring in most tropical forests remain difficult to predict. This uncertainty could potentially exacerbate threats to some of the thousands of timber-valuable tree species in the Amazon. Our objective was to determine to what extent long-term responses of tree species regeneration in logged forests can be explained by their functional traits. We integrate functional trait data for 13 leaf, stem, and seed traits from 25 canopy tree species with a range of life histories, such as the pioneer Goupia glabra and the shade-tolerant Iryanthera hostmannii, together with over 30 yr of sapling monitoring in permanent plots spanning a gradient of harvest intensity at the Paracou Forest Disturbance Experiment (PFDE), French Guiana. We anticipated that more intensive logging would increase recruitment of pioneer species with higher specific leaf area, lower wood densities, and smaller seeds, due to the removal of canopy trees. We define a recruitment response metric to compare sapling regeneration to timber harvest intensity across species. Although not statistically significant, sapling recruitment decreased with logging intensity for eight of 23 species and these species tended to have large seeds and dense wood. A generalized linear mixed model fit using specific leaf area, seed mass, and twig density data explained about 45% of the variability in sapling dynamics. Effects of specific leaf area outweighed those of seed mass and wood density in explaining recruitment dynamics of the sapling community in response to increasing logging intensity. The most intense treatment at the PFDE, which includes stand thinning of non-timber-valuable adult trees and poison-girdling for competitive release, showed evidence of shifting community composition in sapling regeneration at the 30-yr mark, toward species with less dense wood, lighter seeds, and higher specific leaf area. Our results indicate that high-intensity logging can have lasting effects on stand regeneration dynamics and that functional traits can help simplify general trends of sapling recruitment for highly diverse logged tropical forests.

Intraspecific variation in Janzen-Connell effect is mediated by stress and plant-soil feedbacks.

Janzen-Connell (JC) effects, hypothesized to be partially driven by negative plant-soil feedbacks (PSFs), are considered to be a key mechanism that regulates tropical forest plant diversity and coexistence. However, intraspecific variation in JC effects may weaken this mechanism, with the strength of PSFs being a potentially key variable process. We conducted a manipulated experiment with seedlings from two populations of Pometia pinnata (Sapindaceae), a tropical tree species in southwest China. We aimed to measure the intraspecific difference in PSF magnitude caused by inoculating the soil from different P. pinnata source populations and growing seedlings under differing light intensity and water availability treatments, and at varying plant densities. We found negative PSFs for both populations with the inoculum soil originating from the same sites, but PSFs differed significantly with the inoculum soil from different sites. PSF strength responded differently to biotic and abiotic drivers; PSF strength was weaker in low moisture and high light treatments than in high moisture and low light treatments. Our study documents intraspecific variation in JC effects: specifically, P. pinnata have less defenses to their natively-sourced soil, but are more defensive to the soil feedbacks from soil sourced from other populations. Our results imply that drought and light intensity tended to weaken JC effects, which may result in loss of species diversity with climate change.

Survival, growth, and functional traits of tropical wet forest tree seedlings across an experimental soil moisture gradient in Puerto Rico.

Droughts are predicted to become more frequent and intense in many tropical regions, which may cause shifts in plant community composition. Especially in diverse tropical communities, understanding how traits mediate demographic responses to drought can help provide insight into the effects of climate change on these ecosystems. To understand tropical tree responses to reduced soil moisture, we grew seedlings of eight species across an experimental soil moisture gradient at the Luquillo Experimental Forest, Puerto Rico. We quantified survival and growth over an 8-month period and characterized demographic responses in terms of tolerance to low soil moisture-defined as survival and growth rates under low soil moisture conditions-and sensitivity to variation in soil moisture-defined as more pronounced changes in demographic rates across the observed range of soil moisture. We then compared demographic responses with interspecific variation in a suite of 11 (root, stem, and leaf) functional traits, measured on individuals that survived the experiment. Lower soil moisture was associated with reduced survival and growth but traits mediated species-specific responses. Species with relatively conservative traits (e.g., high leaf mass per area), had higher survival at low soil moisture whereas species with more extensive root systems were more sensitive to soil moisture, in that they exhibited more pronounced changes in growth across the experimental soil moisture gradient. Our results suggest that increasing drought will favor species with more conservative traits that confer greater survival in low soil moisture conditions.

Responses of a common tropical epiphyte, Asplenium nidus, to changes in water and nutrient availability.

Epiphytes are highly dependent on atmospheric inputs of water and nutrients. Reductions in water availability associated with warming and climate change and continual atmospheric nitrogen (N) deposition can affect plant growth but few studies have evaluated the effects of changes in both water and nutrient availabilities on epiphytes. We experimentally tested whether epiphyte growth is more water- or nutrient-limited, if nutrient limitation was stronger for nitrogen or phosphorus, and whether nutrient limitation interacts with water availability. We applied watering (high and low) and nutrient addition (control, +N, +P, +N+P) treatments to greenhouse-grown Asplenium nidus, a common epiphytic fern found in many tropical and subtropical wet forests. We measured leaf area production and leaf elemental concentrations to assess how A. nidus growth and physiology respond to changes in water and nutrient availabilities. We found that leaf growth of A. nidus was more affected by water availability than nutrient addition and the effect of adding nutrients was not fully realized under low-water availability. Among the different nutrient treatments, +N+P had the greatest effects on A. nidus growth and physiology in both watering treatments. Watering treatment changed leaf elemental concentrations but not their ratios (i.e. C:N and N:P). Nutrient addition altered C:N and N:P ratios and increased the concentration of the added elements in leaves, with more pronounced increases in the high-watering treatment. We conclude that the growth of A. nidus is more water- than nutrient-limited. When nutrient limitation occurs (i.e. under high-water availability), nutrient co-limitation is stronger than limitation by N or P alone. This result taken together with studies of other epiphytes suggests greater water than nutrient limitation is likely widespread among epiphytic plants. The limited effects of nutrient addition in the low-water treatment suggest that the effect of atmospheric N deposition on epiphyte growth will be limited when water availability is low.

Water levels primarily drive variation in photosynthesis and nutrient use of scrub Red Mangroves in the southeastern Florida Everglades.

We investigated how mangrove-island micro-elevation (i.e., habitat: center vs edge) affects tree physiology in a scrub mangrove forest of the southeastern Everglades. We measured leaf gas exchange rates of scrub Rhizophora mangle L. trees monthly during 2019, hypothesizing that CO2 assimilation (Anet) and stomatal conductance (gsw) would decline with increasing water levels and salinity, expecting more considerable differences at mangrove-island edges than centers, where physiological stress is greatest. Water levels varied between 0 and 60 cm from the soil surface, rising during the wet season (May-October) relative to the dry season (November-April). Porewater salinity ranged from 15 to 30 p.p.t., being higher at mangrove-island edges than centers. Anet maximized at 15.1 µmol m-2 s-1, and gsw was typically <0.2 mol m-2 s-1, both of which were greater in the dry than the wet season and greater at island centers than edges, with seasonal variability being roughly equal to variation between habitats. After accounting for season and habitat, water level positively affected Anet in both seasons but did not affect gsw. Our findings suggest that inundation stress (i.e., water level) is the primary driver of variation in leaf gas exchange rates of scrub mangroves in the Florida Everglades, while also constraining Anet more than gsw. The interaction between inundation stress due to permanent flooding and habitat varies with season as physiological stress is alleviated at higher-elevation mangrove-island center habitats during the dry season. Freshwater inflows during the wet season increase water levels and inundation stress at higher-elevation mangrove-island centers, but also potentially alleviate salt and sulfide stress in soils. Thus, habitat heterogeneity leads to differences in nutrient and water acquisition and use between trees growing in island centers versus edges, creating distinct physiological controls on photosynthesis, which likely affect carbon flux dynamics of scrub mangroves in the Everglades.

A general pattern of trade-offs between ecosystem resistance and resilience to tropical cyclones.

Tropical cyclones drive coastal ecosystem dynamics, and their frequency, intensity, and spatial distribution are predicted to shift with climate change. Patterns of resistance and resilience were synthesized for 4138 ecosystem time series from n = 26 storms occurring between 1985 and 2018 in the Northern Hemisphere to predict how coastal ecosystems will respond to future disturbance regimes. Data were grouped by ecosystems (fresh water, salt water, terrestrial, and wetland) and response categories (biogeochemistry, hydrography, mobile biota, sedentary fauna, and vascular plants). We observed a repeated pattern of trade-offs between resistance and resilience across analyses. These patterns are likely the outcomes of evolutionary adaptation, they conform to disturbance theories, and they indicate that consistent rules may govern ecosystem susceptibility to tropical cyclones.

Tropical Cyclone Ecology: A Scale-Link Perspective.

Tropical cyclones are increasing in intensity and size and, thus, are poised to increase in importance as disturbance agents. Our understanding of cyclone ecology is biased towards the North Atlantic Basin, because cyclone effects do differ across oceanic basins. Cyclones have both short and long-term effects across the levels of biological organization, but we lack a scale-perspective of cyclone ecology. Effects on individual trees, such as defoliation or branch stripping and uprooting, are mechanistically linked to effects at the community and ecosystem levels, including forest productivity and stand regeneration time. Forest dwarfing via the gradual removal of taller trees by cyclones over many generations illustrates that cyclones shape forest structure through the accumulation of short-term effects over longer timescales.

Soil nitrogen concentration mediates the relationship between leguminous trees and neighbor diversity in tropical forests.

Legumes provide an essential service to ecosystems by capturing nitrogen from the atmosphere and delivering it to the soil, where it may then be available to other plants. However, this facilitation by legumes has not been widely studied in global tropical forests. Demographic data from 11 large forest plots (16-60 ha) ranging from 5.25° S to 29.25° N latitude show that within forests, leguminous trees have a larger effect on neighbor diversity than non-legumes. Where soil nitrogen is high, most legume species have higher neighbor diversity than non-legumes. Where soil nitrogen is low, most legumes have lower neighbor diversity than non-legumes. No facilitation effect on neighbor basal area was observed in either high or low soil N conditions. The legume-soil nitrogen positive feedback that promotes tree diversity has both theoretical implications for understanding species coexistence in diverse forests, and practical implications for the utilization of legumes in forest restoration.

Publisher Correction: Open Science principles for accelerating trait-based science across the Tree of Life.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Open Science principles for accelerating trait-based science across the Tree of Life.

Synthesizing trait observations and knowledge across the Tree of Life remains a grand challenge for biodiversity science. Species traits are widely used in ecological and evolutionary science, and new data and methods have proliferated rapidly. Yet accessing and integrating disparate data sources remains a considerable challenge, slowing progress toward a global synthesis to integrate trait data across organisms. Trait science needs a vision for achieving global integration across all organisms. Here, we outline how the adoption of key Open Science principles-open data, open source and open methods-is transforming trait science, increasing transparency, democratizing access and accelerating global synthesis. To enhance widespread adoption of these principles, we introduce the Open Traits Network (OTN), a global, decentralized community welcoming all researchers and institutions pursuing the collaborative goal of standardizing and integrating trait data across organisms. We demonstrate how adherence to Open Science principles is key to the OTN community and outline five activities that can accelerate the synthesis of trait data across the Tree of Life, thereby facilitating rapid advances to address scientific inquiries and environmental issues. Lessons learned along the path to a global synthesis of trait data will provide a framework for addressing similarly complex data science and informatics challenges.

Botanic gardens are an untapped resource for studying the functional ecology of tropical plants.

Functional traits are increasingly used to understand the ecology of plants and to predict their responses to global changes. Unfortunately, trait data are unavailable for the majority of plant species. The lack of trait data is especially prevalent for hard-to-measure traits and for tropical plant species, potentially owing to the many inherent difficulties of working with species in remote, hyperdiverse rainforest systems. The living collections of botanic gardens provide convenient access to large numbers of tropical plant species and can potentially be used to quickly augment trait databases and advance our understanding of species' responses to climate change. In this review, we quantitatively assess the availability of trait data for tropical versus temperate species, the diversity of species available for sampling in several exemplar tropical botanic gardens and the validity of garden-based leaf and root trait measurements. Our analyses support the contention that the living collections of botanic gardens are a valuable scientific resource that can contribute significantly to research on plant functional ecology and conservation.This article is part of the theme issue 'Biological collections for understanding biodiversity in the Anthropocene'.