Individual canopy tree species maps for the National Ecological Observatory Network.

The ecology of forest ecosystems depends on the composition of trees. Capturing fine-grained information on individual trees at broad scales provides a unique perspective on forest ecosystems, forest restoration, and responses to disturbance. Individual tree data at wide extents promises to increase the scale of forest analysis, biogeographic research, and ecosystem monitoring without losing details on individual species composition and abundance. Computer vision using deep neural networks can convert raw sensor data into predictions of individual canopy tree species through labeled data collected by field researchers. Using over 40,000 individual tree stems as training data, we create landscape-level species predictions for over 100 million individual trees across 24 sites in the National Ecological Observatory Network (NEON). Using hierarchical multi-temporal models fine-tuned for each geographic area, we produce open-source data available as 1 km2 shapefiles with individual tree species prediction, as well as crown location, crown area, and height of 81 canopy tree species. Site-specific models had an average performance of 79% accuracy covering an average of 6 species per site, ranging from 3 to 15 species per site. All predictions are openly archived and have been uploaded to Google Earth Engine to benefit the ecology community and overlay with other remote sensing assets. We outline the potential utility and limitations of these data in ecology and computer vision research, as well as strategies for improving predictions using targeted data sampling.

Incorporating plant phenological responses into species distribution models reduces estimates of future species loss and turnover.

Anthropogenetic climate change has caused range shifts among many species. Species distribution models (SDMs) are used to predict how species ranges may change in the future. However, most SDMs rarely consider how climate-sensitive traits, such as phenology, which affect individuals' demography and fitness, may influence species' ranges. Using > 120 000 herbarium specimens representing 360 plant species distributed across the eastern United States, we developed a novel 'phenology-informed' SDM that integrates phenological responses to changing climates. We compared the ranges of each species forecast by the phenology-informed SDM with those from conventional SDMs. We further validated the modeling approach using hindcasting. When examining the range changes of all species, our phenology-informed SDMs forecast less species loss and turnover under climate change than conventional SDMs. These results suggest that dynamic phenological responses of species may help them adjust their ecological niches and persist in their habitats as the climate changes. Plant phenology can modulate species' responses to climate change, mitigating its negative effects on species persistence. Further application of our framework will contribute to a generalized understanding of how traits affect species distributions along environmental gradients and facilitate the use of trait-based SDMs across spatial and taxonomic scales.

Climate-driven, but dynamic and complex? A reconciliation of competing hypotheses for species' distributions.

Estimates of the percentage of species "committed to extinction" by climate change range from 15% to 37%. The question is whether factors other than climate need to be included in models predicting species' range change. We created demographic range models that include climate vs. climate-plus-competition, evaluating their influence on the geographic distribution of Pinus edulis, a pine endemic to the semiarid southwestern U.S. Analyses of data on 23,426 trees in 1941 forest inventory plots support the inclusion of competition in range models. However, climate and competition together only partially explain this species' distribution. Instead, the evidence suggests that climate affects other range-limiting processes, including landscape-scale, spatial processes such as disturbances and antagonistic biotic interactions. Complex effects of climate on species distributions-through indirect effects, interactions, and feedbacks-are likely to cause sudden changes in abundance and distribution that are not predictable from a climate-only perspective.

Addressing bias in faculty retention.

The field of ecology in the United States is not fully harnessing the diverse perspectives of the American population. Two major limitations to advancing diversity and inclusion include insufficient awareness of biased attitudes and the lack of large-scale faculty engagement in diversity and inclusion programs. Academic institutions must recognize and value individuals that participate in diversity and inclusion programs. Valuing this work will motivate all ecologists to accept the responsibility for these efforts and not simply assume that the few minorities in their field can do this work.

Broadening the ecological mindset.

Over the past three decades, the Harvard Forest Summer Research Program in Ecology (HF-SRPE) has been at the forefront of expanding the ecological tent for minoritized or otherwise marginalized students. By broadening the definition of ecology to include fields such as data science, software engineering, and remote sensing, we attract a broader range of students, including those who may not prioritize field experiences or who may feel unsafe working in rural or urban field sites. We also work towards a more resilient society in which minoritized or marginalized students can work safely, in part by building teams of students and mentors. Teams collaborate on projects that require a diversity of approaches and create opportunities for students and mentors alike to support one another and share leadership. Finally, HF-SRPE promotes an expanded view of what it means to become an ecologist. We value and support diverse career paths for ecologists to work in all parts of society, to diversify the face of ecology, and to bring different perspectives together to ensure innovations in environmental problem solving for our planet.

Changes in taxonomic and phylogenetic diversity in the Anthropocene.

To better understand how ecosystems are changing, a multifaceted approach to measuring biodiversity that considers species richness (SR) and evolutionary history across spatial scales is needed. Here, we compiled 162 datasets for fish, bird and plant assemblages across the globe and measured how taxonomic and phylogenetic diversity changed at different spatial scales (within site α diversity and between sites spatial ß diversity). Biodiversity change is measured from these datasets in three ways: across land use gradients, from species lists, and through sampling of the same locations across two time periods. We found that local SR and phylogenetic α diversity (Faith's PD (phylogenetic diversity)) increased for all taxonomic groups. However, when measured with a metric that is independent of SR (phylogenetic species variation, PSV), phylogenetic α diversity declined for all taxonomic groups. Land use datasets showed declines in SR, Faith's PD and PSV. For all taxonomic groups and data types, spatial taxonomic and phylogenetic ß diversity decreased when measured with Sorensen dissimilarity and phylogenetic Sorensen dissimilarity, respectively, providing strong evidence of global biotic homogenization. The decoupling of α and ß diversity, as well as taxonomic and phylogenetic diversity, highlights the need for a broader perspective on contemporary biodiversity changes. Conservation and environmental policy decisions thus need to consider biodiversity beyond local SR to protect biodiversity and ecosystem services.

Towards connecting biodiversity and geodiversity across scales with satellite remote sensing.

ISSUE: Geodiversity (i.e., the variation in Earth's abiotic processes and features) has strong effects on biodiversity patterns. However, major gaps remain in our understanding of how relationships between biodiversity and geodiversity vary over space and time. Biodiversity data are globally sparse and concentrated in particular regions. In contrast, many forms of geodiversity can be measured continuously across the globe with satellite remote sensing. Satellite remote sensing directly measures environmental variables with grain sizes as small as tens of metres and can therefore elucidate biodiversity-geodiversity relationships across scales. EVIDENCE: We show how one important geodiversity variable, elevation, relates to alpha, beta and gamma taxonomic diversity of trees across spatial scales. We use elevation from NASA's Shuttle Radar Topography Mission (SRTM) and c. 16,000 Forest Inventory and Analysis plots to quantify spatial scaling relationships between biodiversity and geodiversity with generalized linear models (for alpha and gamma diversity) and beta regression (for beta diversity) across five spatial grains ranging from 5 to 100 km. We illustrate different relationships depending on the form of diversity; beta and gamma diversity show the strongest relationship with variation in elevation. CONCLUSION: With the onset of climate change, it is more important than ever to examine geodiversity for its potential to foster biodiversity. Widely available satellite remotely sensed geodiversity data offer an important and expanding suite of measurements for understanding and predicting changes in different forms of biodiversity across scales. Interdisciplinary research teams spanning biodiversity, geoscience and remote sensing are well poised to advance understanding of biodiversity-geodiversity relationships across scales and guide the conservation of nature.

Tropical bird species have less variable body sizes.

Ecologists have often predicted that species' niche breadths should decline towards the Equator. Dan Janzen arrived at this prediction based on climatic constraints, while Robert MacArthur argued that a latitudinal gradient in resource specialization drives the pattern. This idea has some support when it comes to thermal niches, but has rarely been explored for other niche dimensions. Body size is linked to niche dimensions related to diet, competition and environmental tolerance in vertebrates. We identified 68 pairs of tropical and nontropical sister bird species using a comprehensive phylogeny and used the VertNet specimen database to ask whether tropical birds have lower intraspecific body-size variation than their nontropical sister species. Our results show that tropical species have less intraspecific variability in body mass ([Formula: see text]; p = 0.009). Variation in body-size variability was poorly explained by both abiotic and biotic drivers; thus the mechanisms underlying the pattern are still unclear. The lower variation in body size of tropical bird species may have evolved in response to more stable climates and resource environments.

Observed forest sensitivity to climate implies large changes in 21st century North American forest growth.

Predicting long-term trends in forest growth requires accurate characterisation of how the relationship between forest productivity and climatic stress varies across climatic regimes. Using a network of over two million tree-ring observations spanning North America and a space-for-time substitution methodology, we forecast climate impacts on future forest growth. We explored differing scenarios of increased water-use efficiency (WUE) due to CO2 -fertilisation, which we simulated as increased effective precipitation. In our forecasts: (1) climate change negatively impacted forest growth rates in the interior west and positively impacted forest growth along the western, southeastern and northeastern coasts; (2) shifting climate sensitivities offset positive effects of warming on high-latitude forests, leaving no evidence for continued 'boreal greening'; and (3) it took a 72% WUE enhancement to compensate for continentally averaged growth declines under RCP 8.5. Our results highlight the importance of locally adapted forest management strategies to handle regional differences in growth responses to climate change.

Seedling survival responses to conspecific density, soil nutrients, and irradiance vary with age in a tropical forest.

Understanding processes that promote species coexistence is integral to diversity maintenance. In hyperdiverse tropical forests, local conspecific density (LCD) and light are influential to woody seedling recruitment and soil nutrients are often limiting, yet the simultaneous effects of these factors on seedling survival across time remain unknown. We fit species- and age-specific models to census and resource data of seedlings of 68 woody species from a Costa Rican wet tropical forest. In decreasing order of prevalence, seedling survivorship was related to LCD, soil base cations, irradiance, nitrogen, and phosphorus. Species-specific responses to factors did not covary, providing evidence that species life history strategies have not converged to one continuum of high-surviving stress tolerant to low-surviving stress intolerant species. Survival responses to all factors varied over the average seedling's lifetime, indicating seedling requirements change with age and conclusions drawn about processes important to species coexistence depend on temporal resolution.

Speeding up ecological and evolutionary computations in R; essentials of high performance computing for biologists.

Computation has become a critical component of research in biology. A risk has emerged that computational and programming challenges may limit research scope, depth, and quality. We review various solutions to common computational efficiency problems in ecological and evolutionary research. Our review pulls together material that is currently scattered across many sources and emphasizes those techniques that are especially effective for typical ecological and environmental problems. We demonstrate how straightforward it can be to write efficient code and implement techniques such as profiling or parallel computing. We supply a newly developed R package (aprof) that helps to identify computational bottlenecks in R code and determine whether optimization can be effective. Our review is complemented by a practical set of examples and detailed Supporting Information material (S1-S3 Texts) that demonstrate large improvements in computational speed (ranging from 10.5 times to 14,000 times faster). By improving computational efficiency, biologists can feasibly solve more complex tasks, ask more ambitious questions, and include more sophisticated analyses in their research.

Plasticity and not adaptation is the primary source of temperature-mediated variation in flowering phenology in North America.

Phenology varies widely over space and time because of its sensitivity to climate. However, whether phenological variation is primarily generated by rapid organismal responses (plasticity) or local adaptation remains unresolved. Here we used 1,038,027 herbarium specimens representing 1,605 species from the continental United States to measure flowering-time sensitivity to temperature over time (Stime) and space (Sspace). By comparing these estimates, we inferred how adaptation and plasticity historically influenced phenology along temperature gradients and how their contributions vary among species with different phenology and native climates and among ecoregions differing in species composition. Parameters Sspace and Stime were positively correlated (r = 0.87), of similar magnitude and more frequently consistent with plasticity than adaptation. Apparent plasticity and adaptation generated earlier flowering in spring, limited responsiveness in late summer and delayed flowering in autumn in response to temperature increases. Nonetheless, ecoregions differed in the relative contributions of adaptation and plasticity, from consistently greater importance of plasticity (for example, southeastern United States plains) to their nearly equal importance throughout the season (for example, Western Sierra Madre Piedmont). Our results support the hypothesis that plasticity is the primary driver of flowering-time variation along temperature gradients, with local adaptation having a widespread but comparatively limited role.

Diversity-stability relationships across organism groups and ecosystem types become decoupled across spatial scales.

The relationship between biodiversity and stability, or its inverse, temporal variability, is multidimensional and complex. Temporal variability in aggregate properties, like total biomass or abundance, is typically lower in communities with higher species diversity (i.e., the diversity-stability relationship [DSR]). At broader spatial extents, regional-scale aggregate variability is also lower with higher regional diversity (in plant systems) and with lower spatial synchrony. However, focusing exclusively on aggregate properties of communities may overlook potentially destabilizing compositional shifts. It is not yet clear how diversity is related to different components of variability across spatial scales, nor whether regional DSRs emerge across a broad range of organisms and ecosystem types. To test these questions, we compiled a large collection of long-term metacommunity data spanning a wide range of taxonomic groups (e.g., birds, fish, plants, invertebrates) and ecosystem types (e.g., deserts, forests, oceans). We applied a newly developed quantitative framework for jointly analyzing aggregate and compositional variability across scales. We quantified DSRs for composition and aggregate variability in local communities and metacommunities. At the local scale, more diverse communities were less variable, but this effect was stronger for aggregate than compositional properties. We found no stabilizing effect of γ-diversity on metacommunity variability, but ß-diversity played a strong role in reducing compositional spatial synchrony, which reduced regional variability. Spatial synchrony differed among taxa, suggesting differences in stabilization by spatial processes. However, metacommunity variability was more strongly driven by local variability than by spatial synchrony. Across a broader range of taxa, our results suggest that high γ-diversity does not consistently stabilize aggregate properties at regional scales without sufficient spatial ß-diversity to reduce spatial synchrony.

Pattern and process of biotic homogenization in the New Pangaea.

Human activities have reorganized the earth's biota resulting in spatially disparate locales becoming more or less similar in species composition over time through the processes of biotic homogenization and biotic differentiation, respectively. Despite mounting evidence suggesting that this process may be widespread in both aquatic and terrestrial systems, past studies have predominantly focused on single taxonomic groups at a single spatial scale. Furthermore, change in pairwise similarity is itself dependent on two distinct processes, spatial turnover in species composition and changes in gradients of species richness. Most past research has failed to disentangle the effect of these two mechanisms on homogenization patterns. Here, we use recent statistical advances and collate a global database of homogenization studies (20 studies, 50 datasets) to provide the first global investigation of the homogenization process across major faunal and floral groups and elucidate the relative role of changes in species richness and turnover. We found evidence of homogenization (change in similarity ranging from -0.02 to 0.09) across nearly all taxonomic groups, spatial extent and grain sizes. Partitioning of change in pairwise similarity shows that overall change in community similarity is driven by changes in species richness. Our results show that biotic homogenization is truly a global phenomenon and put into question many of the ecological mechanisms invoked in previous studies to explain patterns of homogenization.

Consistent stabilizing effects of plant diversity across spatial scales and climatic gradients.

Biodiversity has widely been documented to enhance local community stability but whether such stabilizing effects of biodiversity extend to broader scales remains elusive. Here, we investigated the relationships between biodiversity and community stability in natural plant communities from quadrat (1 m2) to plot (400 m2) and regional (5-214 km2) scales and across broad climatic conditions, using an extensive plant community dataset from the National Ecological Observatory Network. We found that plant diversity provided consistent stabilizing effects on total community abundance across three nested spatial scales and climatic gradients. The strength of the stabilizing effects of biodiversity increased modestly with spatial scale and decreased as precipitation seasonality increased. Our findings illustrate the generality of diversity-stability theory across scales and climatic gradients, which provides a robust framework for understanding ecosystem responses to biodiversity and climate changes.

Experimental and observational evidence of negative conspecific density dependence in temperate ectomycorrhizal trees.

Conspecific negative density dependence (CNDD) promotes tree species diversity by reducing recruitment near conspecific adults due to biotic feedbacks from herbivores, pathogens, or competitors. While this process is well-described in tropical forests, tests of temperate tree species range from strong positive to strong negative density dependence. To explain this, several studies have suggested that tree species traits may help predict the strength and direction of density dependence: for example, ectomycorrhizal-associated tree species typically exhibit either positive or weaker negative conspecific density dependence. More generally, the strength of density dependence may be predictably related to other species-specific ecological attributes such as shade tolerance, or the relative local abundance of a species. To test the strength of density dependence and whether it affects seedling community diversity in a temperate forest, we tracked the survival of seedlings of three ectomycorrhizal-associated species experimentally planted beneath conspecific and heterospecific adults on the Prospect Hill tract of the Harvard Forest, in Massachusetts, USA. Experimental seedling survival was always lower under conspecific adults, which increased seedling community diversity in one of six treatments. We compared these results to evidence of CNDD from observed sapling survival patterns of 28 species over approximately 8 years in an adjacent 35-ha forest plot. We tested whether species-specific estimates of CNDD were associated with mycorrhizal association, shade tolerance, and local abundance. We found evidence of significant, negative conspecific density dependence (CNDD) in 23 of 28 species, and positive conspecific density dependence in two species. Contrary to our expectations, ectomycorrhizal-associated species generally exhibited stronger (e.g., more negative) CNDD than arbuscular mycorrhizal-associated species. CNDD was also stronger in more shade-tolerant species but was not associated with local abundance. Conspecific adult trees often have a negative influence on seedling survival in temperate forests, particularly for tree species with certain traits. Here we found strong experimental and observational evidence that ectomycorrhizal-associating species consistently exhibit CNDD. Moreover, similarities in the relative strength of density dependence from experiments and observations of sapling mortality suggest a mechanistic link between negative effects of conspecific adults on seedling and sapling survival and local tree species distributions.

Long-term ecological research and the COVID-19 anthropause: A window to understanding social-ecological disturbance.

The period of disrupted human activity caused by the COVID-19 pandemic, coined the "anthropause," altered the nature of interactions between humans and ecosystems. It is uncertain how the anthropause has changed ecosystem states, functions, and feedback to human systems through shifts in ecosystem services. Here, we used an existing disturbance framework to propose new investigation pathways for coordinated studies of distributed, long-term social-ecological research to capture effects of the anthropause. Although it is still too early to comprehensively evaluate effects due to pandemic-related delays in data availability and ecological response lags, we detail three case studies that show how long-term data can be used to document and interpret changes in air and water quality and wildlife populations and behavior coinciding with the anthropause. These early findings may guide interpretations of effects of the anthropause as it interacts with other ongoing environmental changes in the future, particularly highlighting the importance of long-term data in separating disturbance impacts from natural variation and long-term trends. Effects of this global disturbance have local to global effects on ecosystems with feedback to social systems that may be detectable at spatial scales captured by nationally to globally distributed research networks.

The geodiv r package: Tools for calculating gradient surface metrics.

The geodiv r package calculates gradient surface metrics from imagery and other gridded datasets to provide continuous measures of landscape heterogeneity for landscape pattern analysis. geodiv is the first open-source, command line toolbox for calculating many gradient surface metrics and easily integrates parallel computing for applications with large images or rasters (e.g. remotely sensed data). All functions may be applied either globally to derive a single metric for an entire image or locally to create a texture image over moving windows of a user-defined extent.We present a comprehensive description of the functions available through geodiv. A supplemental vignette provides an example application of geodiv to the fields of landscape ecology and biogeography. geodiv allows users to easily retrieve estimates of spatial heterogeneity for a variety of purposes, enhancing our understanding of how environmental structure influences ecosystem processes. The package works with any continuous imagery and may be widely applied in many fields where estimates of surface complexity are useful.