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Table Mountain pine (Pinus pungens Lamb.) is an imperiled tree species endemic to the southern and central Appalachian Mountains. Generally reliant on fire for regeneration, its fragmented but widespread distribution has declined in recent decades. We quantified the genetic diversity of 26 populations across the range of the species using data from seven highly polymorphic simple sequence repeat (SSR) loci. The species was relatively inbred whereas differentiation among populations was relatively low. Differentiation was significantly but weakly associated with geographic distance among populations. We detected minor genetic differences between northern and southern seed collection zones established based on climate similarity. We conducted a series of simulations using SSR data from 498 seedlings, grown from seed collected from five natural stands of Table Mountain pine, to assess the genetic consequences of different strategies for deploying collected seed in ex situ conservation plantings. Results indicated that reducing the number of families in a planting would not substantially affect the conservation of common alleles but would affect the representation of rare alleles and overall allelic richness. These findings add to our limited knowledge of genetic variation across the distribution of this rare conifer and offer some guidance for its effective genetic conservation.

Thermal sprayed metal coatings are used in many industrial applications, and characterizing the structure and performance of these materials is vital to understanding their behavior in the field. X-ray computed tomography (CT) enables volumetric, nondestructive imaging of these materials, but precise segmentation of this grayscale image data into discrete material phases is necessary to calculate quantities of interest related to material structure. In this work, we present a methodology to automate the CT segmentation process as well as quantify uncertainty in segmentations via deep learning. Neural networks (NNs) have been shown to excel at segmentation tasks; however, memory constraints, class imbalance, and lack of sufficient training data often prohibit their deployment in high resolution volumetric domains. Our 3D convolutional NN implementation mitigates these challenges and accurately segments full resolution CT scans of thermal sprayed materials with maps of uncertainty that conservatively bound the predicted geometry. These bounds are propagated through calculations of material properties such as porosity that may provide an understanding of anticipated behavior in the field.

Biodiversity underlies ecosystem resilience, ecosystem function, sustainable economies, and human well-being. Understanding how biodiversity sustains ecosystems under anthropogenic stressors and global environmental change will require new ways of deriving and applying biodiversity data. A major challenge is that biodiversity data and knowledge are scattered, biased, collected with numerous methods, and stored in inconsistent ways. The Group on Earth Observations Biodiversity Observation Network (GEO BON) has developed the Essential Biodiversity Variables (EBVs) as fundamental metrics to help aggregate, harmonize, and interpret biodiversity observation data from diverse sources. Mapping and analyzing EBVs can help to evaluate how aspects of biodiversity are distributed geographically and how they change over time. EBVs are also intended to serve as inputs and validation to forecast the status and trends of biodiversity, and to support policy and decision making. Here, we assess the feasibility of implementing Genetic Composition EBVs (Genetic EBVs), which are metrics of within-species genetic variation. We review and bring together numerous areas of the field of genetics and evaluate how each contributes to global and regional genetic biodiversity monitoring with respect to theory, sampling logistics, metadata, archiving, data aggregation, modeling, and technological advances. We propose four Genetic EBVs: (i) Genetic Diversity; (ii) Genetic Differentiation; (iii) Inbreeding; and (iv) Effective Population Size (Ne ). We rank Genetic EBVs according to their relevance, sensitivity to change, generalizability, scalability, feasibility and data availability. We outline the workflow for generating genetic data underlying the Genetic EBVs, and review advances and needs in archiving genetic composition data and metadata. We discuss how Genetic EBVs can be operationalized by visualizing EBVs in space and time across species and by forecasting Genetic EBVs beyond current observations using various modeling approaches. Our review then explores challenges of aggregation, standardization, and costs of operationalizing the Genetic EBVs, as well as future directions and opportunities to maximize their uptake globally in research and policy. The collection, annotation, and availability of genetic data has made major advances in the past decade, each of which contributes to the practical and standardized framework for large-scale genetic observation reporting. Rapid advances in DNA sequencing technology present new opportunities, but also challenges for operationalizing Genetic EBVs for biodiversity monitoring regionally and globally. With these advances, genetic composition monitoring is starting to be integrated into global conservation policy, which can help support the foundation of all biodiversity and species' long-term persistence in the face of environmental change. We conclude with a summary of concrete steps for researchers and policy makers for advancing operationalization of Genetic EBVs. The technical and analytical foundations of Genetic EBVs are well developed, and conservation practitioners should anticipate their increasing application as efforts emerge to scale up genetic biodiversity monitoring regionally and globally. ispartof: BIOLOGICAL REVIEWS vol:97 issue:4 pages:1511-1538 ispartof: location:England status: published

AbstractReforestation and restoration efforts have traditionally relied on “local” seed sources as planting stock. The term “local” has different meanings in different locales, since no single set of seed-collection zones has yet been widely adopted across the eastern United States. Given concerns about mitigating the effects of climate change, forest managers are increasingly seeking to move seed sources in a process called assisted migration, which would be facilitated if a common set of seed-collection zones were available. We developed a map of 245 seed-collection zones for 37 states by combining two ecologically important layers: plant hardiness zones and ecoregions. These new seed-collection zones should be used by seed collectors and nurseries to describe the origin of seed for plants and trees in the eastern United States. Common garden studies remain the gold standard for determining how far to move seed from any particular seed-collection zone. For species that have not been field-tested, seed movement that is limited either within a seed-collection zone or between adjacent seed-collection zones is a reasonable general guideline.

The mitochondrial phylogeography of some conifers shows evidence of introgression from sympatric congeners, with mitochondrial lineages not always reflecting species. This suggests that unique mitochondrial haplotypes previously reported in the ponderosa pines (Pinus subsection Ponderosae) from the USA might be more widespread in taxa not yet sampled. Recent nuclear and plastome phylogenies placed Pinus ponderosa paraphyletic in relation to Ponderosae in Mexico and Central America and confirmed that sympatric Pinus jeffreyi is more closely related to the California big-cone pines (Pinus subsection Sabinianae). We describe a broad survey of the repeated motifs in nad1 intron 2 of Ponderosae and Sabinianae, which revealed that most of the 27 mitochondrial haplotypes were not exclusive to a taxon but showed strong geographic patterns. In surprising contrast to nuclear and plastid phylogenies that resolve a monophyletic P. jeffreyi, unidirectional mitochondrial capture by P. jeffreyi (Sabinianae) from P. ponderosa was observed in all 28 samples of Jeffrey pine. Confirming the paraphyly of P. ponderosa sensu lato, mitochondrial haplotypes found mostly west and those found mostly east of the Great Basin each have more similarity to haplotypes found in Mexican taxa than they have to each other. Two distinctive haplotypes that were terminal nodes on the network were confirmed to be endemic to the Great Basin, USA, suggesting that they arose in place and have been maintained in isolation. Altogether, our results indicate a history of complex and intriguing mitochondrial relationships among the ponderosa pine species, especially between P. ponderosa and P. jeffreyi.

The annual national report of the Forest Health Monitoring (FHM) program of the Forest Service, U.S. Department of Agriculture, presents forest health status and trends from a national or multi-State regional perspective using a variety of sources, introduces new techniques for analyzing forest health data, and summarizes results of recently completed Evaluation Monitoring projects funded through the FHM national program. In this 20th edition in a series of annual reports, national survey data are used to identify recent geographic patterns of insect and disease activity. Satellite data are employed to detect geographic patterns of forest fire occurrence. Recent drought and moisture surplus conditions are compared across the conterminous United States. Data collected by the Forest Inventory and Analysis (FIA) program are employed to detect regional differences in tree mortality. Nearly 20 years of FIA coarse woody material (CWM) data are examined across spatial scales to assess change over time in CWM biomass, diameter and decay class, and species composition. Case studies demonstrate the potential of rFIA, an open-source R package developed to improve the accessibility of FIA data and offer enhanced flexibility in estimation strategies, to advance forest health evaluation and monitoring. Six recently completed Evaluation Monitoring projects are summarized, addressing forest health concerns at smaller scales.

International agreements such as the Convention on Biological Diversity (CBD) have committed to conserve, and sustainably and equitably use, biodiversity. The CBD is a vital instrument for global conservation because it guides 195 countries and the European Union in setting priorities and allocating resources, and requires regular reporting on progress. However, the CBD and similar policy agreements have often neglected genetic diversity. This is a critical gap because genetic diversity underlies adaptation to environmental change and ecosystem resilience. Here we aim to inform future policy, monitoring, and reporting efforts focused on limiting biodiversity loss by conducting the largest yet evaluation of how Parties to the CBD report on genetic diversity. A large, globally representative sample of 114 CBD National Reports was examined to assess reported actions, progress, values and indicators related to genetic diversity. Although the importance of genetic diversity is recognized by most Parties to the CBD, genetic diversity targets mainly addressed variation within crops and livestock (a small fraction of all species). Reported actions to conserve genetic diversity primarily concerned ex situ facilities and legislation, rather than monitoring and in situ intervention. The most commonly reported status indicators are not well correlated to maintaining genetic diversity. Lastly, few reports mentioned genetic monitoring using DNA data, indigenous use and knowledge of genetic diversity, or development of strategies to conserve genetic diversity. We make several recommendations for the post-2020 CBD Biodiversity Framework, and similar efforts such as IPBES, to improve awareness, assessment, and monitoring of genetic diversity, and facilitate consistent and complete reporting in the future.

Ecological communities often exhibit greater resistance to biological invasions when these communities consist of species that are not closely related. The effective size of this resistance, however, varies geographically. Here we investigate the drivers of this heterogeneity in the context of known contributions of native trees to the resistance of forests in the eastern United States of America to plant invasions. Using 42,626 spatially referenced forest community observations, we quantified spatial heterogeneity in relationships between evolutionary relatedness amongst native trees and both invasive plant species richness and cover. We then modelled the variability amongst the 91 ecological sections of our study area in the slopes of these relationships in response to three factors known to affect invasion and evolutionary relationships –environmental harshness (as estimated via tree height), relative tree density and environmental variability. Invasive species richness and cover declined in plots having less evolutionarily related native trees. The degree to which they did, however, varied considerably amongst ecological sections. This variability was explained by an ecological section’s mean maximum tree height and, to a lesser degree, SD in maximum tree height (R2GLMM = 0.47 to 0.63). In general, less evolutionarily related native tree communities better resisted overall plant invasions in less harsh forests and in forests where the degree of harshness was more homogenous. These findings can guide future investigations aimed at identifying the mechanisms by which evolutionary relatedness of native species affects exotic species invasions and the environmental conditions under which these effects are most pronounced.

United States forestland is an important ecosystem type, land cover, land use, and economic resource that is facing several drivers of change including climatic. Because of its significance, forestland was identified through the National Climate Assessment (NCA) as a key sector and system of concern to be included in a system of climate indicators as part of a sustained assessment effort. Here, we describe 11 informative core indicators of forests and climate change impacts with metrics available or nearly available for use in the NCA efforts. The recommended indicators are based on a comprehensive conceptual model which recognizes forests as a land use, an ecosystem, and an economic sector. The indicators cover major forest attributes such as extent, structural components such as biomass, functions such as growth and productivity, and ecosystem services such as biodiversity and outdoor recreation. Interactions between humans and forests are represented through indicators focused on the wildland-urban interface, cost to mitigate wildfire risk, and energy produced from forest-based biomass. Selected indicators also include drought and disturbance from both wildfires and biotic agents. The forest indicators presented are an initial set that will need further refinement in coordination with other NCA indicator teams. Our effort ideally will initiate the collection of critical measurements and observations and lead to additional research on forest-climate indicators.

Genetic diversity is critically important for all species-domesticated and wild- to adapt to environmental change, and for ecosystem resilience to extreme events. International agreements such as the Convention on Biological Diversity (CBD) have committed to conserve and sustainably and equitably use all levels of biodiversity-genes, species and ecosystems-globally. However, assessment and monitoring of genetic diversity are often overlooked, and there are large knowledge and policy gaps regarding genetic diversity conservation. In this study, we present the first quantitative analysis of genetic diversity assessments conducted by Parties to the CBD. We conducted a detailed, systematic analysis of 114 CBD 5th (submitted 2014) and 6th (submitted 2018) National Reports to quantitatively assess actions, progress on targets, values and indicators related to genetic diversity. First, we found that the importance of genetic diversity is recognised by most Parties to the CBD, and that recognition increased over time. However, genetic targets mainly addressed genetic diversity within cultivated plants, farm animals, and crop wild relatives, with little focus on other wild species. Also, actions for conserving genetic diversity primarily concerned ex-situ facilities and policy, rather than monitoring and intervention for maintaining genetic diversity in situ. The most commonly used indicators of genetic diversity status were the number of genetic resources in conservation facilities, number of threatened breeds, and Red List Index, which are not well correlated to genetic erosion in most species -- highlighting that genetic change is poorly monitored by current indicators. Lastly, analyses of genetic data observations, indigenous use and knowledge of genetic diversity, and strategies being developed and implemented to conserve genetic diversity are highly under-reported. We make several recommendations for the post-2020 CBD Biodiversity Framework to improve awareness, assessment, and monitoring, and facilitate consistent and complete reporting of progress of genetic diversity in future National Reports.Article Impact StatementAn analysis of genetic diversity in CBD National Reports neglects non-domesticated species and demonstrates need for sufficient indicators.

Because forest tree species face serious threats including insect and disease epidemics, climate change, and forest fragmentation and conversion, prioritizing species and forests for conservation is an essential management goal. This paper describes a species prioritization approach that incorporates both the rarity of species, because of the increased vulnerability associated with rare species, and their evolutionary distinctiveness (ED), a measure of evolutionary originality. Rarity and ED scores, and scores for the two combined, were calculated for 352 North American forest tree species. A weak but significant phylogenetic signal was associated with species rarity. The scores were used to weight species importance values on approximately 130,000 forest inventory plots across the conterminous United States. The resulting plot-level estimates of conservation value were employed to identify geographic hotspots of forests with high conservation value, and to assess whether forests with protected status effectively conserve rarity and ED. Rarity hotspots were detected in California, the Southwest, central Texas, and Florida. Hotspots of ED included locations along the Pacific Coast, in the Northern Rockies, and in scattered eastern locations. Protected forest areas across the United States effectively conserve ED, but not rarity. In fact, rarity was lowest in areas with the highest protection, and highest in areas with no or unknown protected status. Multiple-use protected areas had higher ED, but not rarity, than restricted-use protected areas. Protected area effectiveness varied across the country. Such spatially explicit assessment approaches can help determine which forests to target for monitoring efforts and pro-active management activities.

Unique responses to climate change can occur across intraspecific levels, resulting in individualistic adaptation or movement patterns among populations within a given species. Thus, the need to model potential responses among genetically distinct populations within a species is increasingly recognized. However, predictive models of future distributions are regularly fit at the species level, often because intraspecific variation is unknown or is identified only within limited sample locations. In this study, we considered the role of intraspecific variation to shape the geographic distribution of ponderosa pine (Pinus ponderosa), an ecologically and economically important tree species in North America. Morphological and genetic variation across the distribution of ponderosa pine suggest the need to model intraspecific populations: the two varieties (var. ponderosa and var. scopulorum) and several haplotype groups within each variety have been shown to occupy unique climatic niches, suggesting populations have distinct evolutionary lineages adapted to different environmental conditions. We utilized a recently available, geographically widespread dataset of intraspecific variation (haplotypes) for ponderosa pine and a recently devised lineage distance modeling approach to derive additional, likely intraspecific occurrence locations. We confirmed the relative uniqueness of each haplotype-climate relationship using a niche-overlap analysis, and developed ecological niche models (ENMs) to project the distribution for two varieties and eight haplotypes under future climate forecasts. Future projections of haplotype niche distributions generally revealed greater potential range loss than predicted for the varieties. This difference may reflect intraspecific responses of distinct evolutionary lineages. However, directional trends are generally consistent across intraspecific levels, and include a loss of distributional area and an upward shift in elevation. Our results demonstrate the utility in modeling intraspecific response to changing climate and they inform management and conservation strategies, by identifying haplotypes and geographic areas that may be most at risk, or most secure, under projected climate change.

Eastern hemlock (Tsuga canadensis) occupies a large swath of eastern North America and has historically undergone range expansion and contraction resulting in several genetically separate lineages. This conifer is currently experiencing mortality across most of its range following infestation of a non-native insect. With the goal of better understanding the current and future conservation potential of the species, we evaluate ecological differences among populations within these genetically defined clusters, which were previously inferred using nuclear microsatellite molecular markers from 58 eastern hemlock populations. We sub-divide these clusters into four genetic zones to differentiate putative north-central, north-east and southeast (SE) and southwest evolutionary lineages in eastern hemlock. We use demographic data (relative abundance, mortality, and seedling regeneration) from the Forest Inventory Analysis program in conjunction with environmental data to model how these lineages respond to current and future climatic gradients. Ecologically meaningful relationships are explored in the intraspecific context of hemlock abundance distribution and then related to genetic variation. We also assess hemlock's colonization likelihood via a long distance dispersal model and explore its future genetic and ecological conservation potential by combining the future suitable habitats with colonization likelihoods. Results show that future habitats under climate change will markedly

Genetic diversity provides the essential basis for the adaptation and resilience of tree species to environmental stress and change. The genetic conservation of tree species is an urgent global necessity as forest conversion and fragmentation continue apace, damaging insects and pathogens are transported between continents, and climate change alters local habitat suitability. Effective and efficient genetic conservation of tree species presents a substantial challenge because of the lack of basic information about many species, inadequate resources, and a historical lack of coordination within and between conservation sectors. Several cooperative efforts are already under way and are achieving conservation success, but much work remains. The Gene Conservation of Tree Species—Banking on the Future workshop in 2016 enabled the exchange of information and the creation of collaborations among tree conservation stakeholders. Several key themes emerged during the meeting’s presentations and dialogue, which are further explored in this paper. In situ conservation of species is the long-term goal and is often the most efficient approach for preserving the genetic diversity of many forest tree species. Whether existing reserves adequately protect species and are sufficient for future conservation needs is uncertain. Ex situ conservation is an important complement to in situ efforts, acting as an insurance measure against extinction, providing material for restoration, enabling additional research opportunities, and educating the public. Networks of botanic gardens, government agencies, and non-governmental organizations must continue to coordinate ex situ and in situ efforts to improve the efficiency and effectiveness of tree conservation efforts. Assessing and prioritizing which species and populations require genetic conservation and prioritizing among them is a critical need. Two key tree restoration needs are for wider dissemination of planting stock, particularly stock with resistance to insects and pathogens, and for specific silvicultural prescriptions that facilitate restoration efforts. Effective genetic conservation of forest trees will require ongoing cooperation among widely diverse groups of scientists, managers, and policymakers from the public and private sectors.

Hemlock woolly adelgid (Adelges tsugae Annand) (HWA) is an invasive forest insect sweeping across the native range of eastern (Tsuga canadensis [L.] Carr.) and Carolina (Tsuga caroliniana Engelm.) hemlocks, threatening to severely reduce eastern hemlock extent and to push Carolina hemlock to extirpation. HWA poses a significant threat to these eastern US natives, now infesting hemlocks across 19 states and more than 400 counties. For the long-term preservation of the species, ex situ genetic resource conservation efforts such as seed collection, storage, and adelgid-resistant hemlock breeding may all be necessary. To ensure the efficient and effective application of these efforts, it will be necessary to prioritize locations within the native ranges, because it is logistically impossible to apply these efforts to all populations. To build upon 12 years of seed banking for eastern and Carolina hemlock, we applied a novel approach for incorporating multiple dissimilar data sets into a geographic prioritization of areas for the most effective and efficient conservation of genetic diversity. The approach involves integration of geographic information systems with the multi-attribute frontier mapping technique to identify locations across the ranges of these two imperiled species most in need of conservation actions. Specifically, our assessment incorporated four genetic diversity parameters, a climate component, a measure of population disjunctiveness, a measure of local hemlock abundance, and seed collection density to prioritize areas of eastern and Carolina hemlock occurrence for ex situ gene conservation. For each species, the result was a mapped index of locations prioritized by the combined significance of these factors. For eastern hemlock, this assessment assigned the highest priority to disjunct populations and to some areas within the main-body range. With Carolina hemlock, disjunct northern populations and central main-body locations received the highest prioritization. Our prioritization approach could be applied similarly to other species facing pressure from invasive pests or other environmental threats.

Climate change is one of several threats that will increase the likelihood that forest tree species could experience population-level extirpation or species-level extinction. Scientists and managers from throughout the United States Forest Service have cooperated to develop a framework for conservation priority-setting assessments of forest tree species. This framework uses trait data and predictions of expected climate change pressure to categorize and prioritize 339 native tree species for conservation, monitoring, management and restoration across all forested lands in the contiguous United States and Alaska. The framework allows for the quantitative grouping of species into vulnerability classes that may require different management and conservation strategies for maintaining the adaptive genetic variation of the species within each group. This categorization is based on risk factors relating to the species’ (1) exposure to climate change, (2) sensitivity to climate change, and (3) capacity to adapt to climate change. We used K-means clustering to group species into seven classes based on these three vulnerability dimensions. The most vulnerable class encompassed 35 species with high scores for all three vulnerability dimensions. These will require the most immediate conservation intervention. A group of 43 species had high exposure and sensitivity, probably requiring conservation assistance, while a group of 69 species had high exposure and low adaptive capacity, probably needing close monitoring. This assessment tool should be valuable for scientists and managers determining which species and populations to target for monitoring efforts and for pro-active gene conservation and management activities.

Forest understories across the northern United States (US) are a complex of tree seedlings, endemic forbs, herbs, shrubs, and introduced plant species within a forest structure defined by tree and forest floor attributes. The substantial increase in white-tailed deer (Odocoileus virginianus Zimmerman) populations over the past decades has resulted in heavy browse pressure in many of these forests. To gain an objective assessment of the role of deer in forested ecosystems, a region-wide forest inventory across the northern US was examined in concert with white-tailed deer density information compiled at broad scales. Results indicate that deer density may be an additional driver of tree seedling abundance when analyzed along with stand attributes such as aboveground biomass, relative density, and stand age. Tree seedling abundance generally decreased as deer density increased above 5.8 deer km2 for all forest type groups with the exception of oak-dominated forests. Findings indicate that introduced plant species, of which 393 were recorded in this study, increased in areas with higher deer density. The abundance of whitetailed deer is just as important as forest stand and site attributes in the development of forest understories. Given the complexity of forest and land use dynamics across the northern US, this study provides directions for future research as more data linking forest-dependent wildlife and forest dynamics at regional and national scales become available.

Premise of the study Molecular genetic evidence can help delineate taxa in species complexes that lack diagnostic morphological characters. Pinus ponderosa (Pinaceae; subsection Ponderosae) is recognized as a problematic taxon: plastid phylogenies of exemplars were paraphyletic, and mitochondrial phylogeography suggested at least four subdivisions of P. ponderosa. These patterns have not been examined in the context of other Ponderosae species. We hypothesized that putative intraspecific subdivisions might each represent a separate taxon. Methods We genotyped six highly variable plastid simple sequence repeats in 1903 individuals from 88 populations of P. ponderosa and related Ponderosae (P. arizonica, P. engelmannii, and P. jeffreyi). We used multilocus haplotype networks and discriminant analysis of principal components to test clustering of individuals into genetically and geographically meaningful taxonomic units. Key results There are at least four distinct plastid clusters within P. ponderosa that roughly correspond to the geographic distribution of mitochondrial haplotypes. Some geographic regions have intermixed plastid lineages, and some mitochondrial and plastid boundaries do not coincide. Based on relative distances to other species of Ponderosae, these clusters diagnose four distinct taxa. Conclusions Newly revealed geographic boundaries of four distinct taxa (P. benthamiana, P. brachyptera, P. scopulorum, and a narrowed concept of P. ponderosa) do not correspond completely with taxonomies. Further research is needed to understand their morphological and nuclear genetic makeup, but we suggest that resurrecting originally published species names would more appropriately reflect the taxonomy of this checkered classification than their current treatment as varieties of P. ponderosa.

Biological invasions are among the greatest global and regional threats to biomes in the Anthropocene. Islands, in particular, have been perceived to have higher vulnerability to invasions. Because of the dynamic nature of ongoing invasions, distinguishing regional patterns from global patterns and their underlying determinants remains a challenge. We aim to comparatively examine global versus regional patterns of plant invasions and the possible underlying mechanisms. We also test whether there is a difference in degree of invasion and invasibility between mainland areas and islands. We compiled and analyzed data from published sources for 100 mainland areas (i.e., regions, countries, states, and provinces) and 89 islands across the globe. We find that (1) the pool of exotic species available intrinsically decreases as area of the land considered increases (at global scale, all is native), thus global invasion patterns assessed by exotic fraction (proportion of exotics) are primarily determined by land area; (2) because “exotic” is defined relative to the borders of the target region, “boundary effects” can result in regional differences in invasion patterns without any ecological processes being involved; and (3) human population density is closely linked to exotic fraction within regions that are defined by a single administrative border. There were clear differences between global and regional patterns of plant invasions. We observed no difference in the exotic fraction-area relationship between mainland areas and islands, supporting what we refer to as the “island-mainland continuum concept” (i.e., no clear separation in the degree of invasion between islands and mainland area with regard to the effects of area). Because of scale-dependency in many observed patterns, future focus should be placed on the links between local, regional, and global invasion patterns.

Despite conservation efforts, most forest ecosystems worldwide are affected by biotic invasions; however, the specific impacts vary across different geographic regions and forest types. The relative contributions of the main drivers such as propagule pressure (e.g., due to human population, travel, and trade), climate, land use, and habitat invasibility remain uncertain. The special issue “Impacts of Nonnative Species on the Health of Natural and Planted Forests” was organized to facilitate timely communications among scientists and managers in different regions and to assist in attempts to improve forest health and maintain long-term sustainability. The special issue addresses broad issues related to forest invasions, including the impacts of nonnative species in various forest ecosystems (e.g., natural vs. urban) and the contributions of land use (e.g., fragmentation), human activity, and climate change to invasion. The new findings include identifying hotspots of potential invasion impacts and their causes, which can help inform policy makers as they develop effective strategies for prevention, early detection or eradication, and forest management.

Research Highlights: We demonstrate a macroscale framework combining an invasibility model with forest inventory data, and evaluate regional forest exposure to harmful invasive plants under different types of forest protection. Background and Objectives: Protected areas are a fundamental component of natural resource conservation. The exposure of protected forests to invasive plants can impede achievement of conservation goals, and the effectiveness of protection for limiting forest invasions is uncertain. We conducted a macroscale assessment of the exposure of protected and unprotected forests to harmful invasive plants in the eastern United States. Materials and Methods: Invasibility (the probability that a forest site has been invaded) was estimated for 82,506 inventory plots from site and landscape attributes. The invaded forest area was estimated by using the inventory sample design to scale up plot invasibility estimates to all forest area. We compared the invasibility and the invaded forest area of seven categories of protection with that of de facto protected (publicly owned) forest and unprotected forest in 13 ecological provinces. Results: We estimate approximately 51% of the total forest area has been exposed to harmful invasive plants, including 30% of the protected forest, 38% of the de facto protected forest, and 56% of the unprotected forest. Based on cumulative invasibility, the relative exposure of protection categories depended on the assumed invasibility threshold. Based on the invaded forest area, the five least-exposed protection categories were wilderness area (13% invaded), national park (18%), sustainable use (26%), nature reserve (31%), and de facto protected Federal land (36%). Of the total uninvaded forest area, only 15% was protected and 14% had de facto protection. Conclusions: Any protection is better than none, and public ownership alone is as effective as some types of formal protection. Since most of the remaining uninvaded forest area is unprotected, landscape-level management strategies will provide the most opportunities to conserve it.

Invader traits (including plant growth form) may play an important, and perhaps overlooked, role in determining macroscale patterns of biological invasions and therefore warrant greater consideration in future investigations aimed at understanding these patterns. To assess this need, we used empirical data from a national-level survey of forest in the contiguous 48 states of the USA to identify geographic hotspots of forest plant invasion for three distinct invasion characteristics: invasive species richness, trait richness (defined as the number of the five following plant growth forms represented by the invasive plants present at a given location: forbs, grasses, shrubs, trees, and vines), and species richness within each growth form. Three key findings emerged. 1) The hotspots identified encompassed from 9 to 23% of the total area of our study region, thereby revealing many forests to be not only invaded, but highly invaded. 2) Substantial spatial disagreement among hotspots of invasive species richness, invasive trait richness, and species richness of invasive plants within each growth form revealed many locations to be hotspots for invader traits, or for particular growth forms of invasive plants, rather than for invasive plants in general. 3) Despite eastern forests exhibiting higher levels of plant invasion than western forests, species richness for invasive forbs and grasses in the west were respectively greater than and equivalent to levels found in the east. Contrasting patterns between eastern and western forests in the number of invasive species detected for each growth form combined with the spatial disagreement found among hotspot types suggests trait-based variability invasion drivers. Our findings reveal invader traits to be an important contributor to macroscale invasion patterns.

As the global biomes are increasingly threatened by human activities, understanding of macroscale patterns and processes is pressingly needed for effective management and policy making. Macrosystems ecology, which studies multiscale eco- logical patterns and processes, has gained growing interest in the research community. However, as a relatively new field in ecology, research in macrosys- tems ecology is facing various challenges. In this special issue, we highlight the following two latest exciting developments in this thriving field: (1) novel tools and methods and (2) new understandings on macroscale patterns and processes. While we believe that the contributions featured in this issue provide promising advancements in macrosystems ecology, we also see multiple challenges for future research including (1) multidisciplinary approaches for long- term and multiscale studies and (2) scaling local patterns and processes to broader scales.

Detecting biotic resistance to biological invasions across large geographic areas may require acknowledging multiple metrics of niche usage and potential spatial heterogeneity in associations between invasive and native species diversity and dominance. Determine (1) if native communities are resistant to biological invasions at macroscales; (2) the metrics that best quantify biotic resistance at these scales; and (3) the degree to which the direction and magnitude of invader-native associations vary with scale and/or location. Using a mixed-effects modeling framework to account for potential sub-regional and cross-scale variability in invader-native associations, we modeled the species richness and cover of invasive plants in 42,626 plots located throughout Eastern USA forests in relationship to plot-level estimates of native tree biomass, species richness, and evolutionary diversity. We found (1) native tree biomass and evolutionary diversity, but not species richness, to be negatively associated with invader establishment and dominance, and thus indicative of biotic resistance; (2) evidence that evolutionary diversity limits invader dominance more than it does invader establishment; (3) evidence of greater invasion resistance in parts of the agriculturally-dominated Midwest and in and around the more-contiguous forests of the Appalachian Mountains; and (4) the magnitude to which native tree biomass and evolutionary diversity limit invasion varies across the ranges of these metrics. These findings illustrate the improved understanding of biotic resistance to invasions that is gained by accounting for sub-regional variability in ecological processes, and underscores the need to determine the factors leading to spatial heterogeneity in biotic resistance.

Fine-scale ecological data collected across broad regions are becoming increasingly available. Appropriate geographic analyses of these data can help identify locations of ecological concern. We present one such approach, spatial association of scalable hexagons (SASH), which identifies locations where ecological phenomena occur at greater or lower frequencies than expected by chance. This approach is based on a sampling frame optimized for spatial neighborhood analysis, adjustable to the appropriate spatial resolution, and applicable to multiple data types. We divided portions of the United States into scalable equal-area hexagonal cells and, using three types of data (field surveys, aerial surveys, satellite imagery), identified geographic clusters of forested areas having high and low values for (1) invasive plant diversity and cover, (2) mountain pine beetle-induced tree mortality, and (3) wildland forest fire occurrences. Using the SASH approach, we detected statistically significant patterns of plant invasion, bark beetle-induced tree mortality, and fire occurrence density that will be useful for understanding macroscale patterns and processes associated with each forest health threat, for assessing its ecological and economic impacts, and for identifying areas where specific management activities may be needed. The presented method is a “big data” analysis tool with potential application for macrosystems ecology studies that require rigorous testing of hypotheses within a spatial framework. This method is a standard component of annual national reports on forest health status and trends across the United States and can be applied easily to other regions and datasets.

Habitat invasibility is a central focus of invasion biology, with implications for basic ecological patterns and processes and for effective invasion management. "Invasibility" is, however, one of the most elusive metrics and misused terms in ecology. Empirical studies and meta-analyses of invasibility have produced inconsistent and even conflicting results. This lack of consistency, and subsequent difficulty in making broad cross-habitat comparisons, stem in part from (1) the indiscriminant use of a closely related, but fundamentally different concept, that of degree of invasion (DI) or level of invasion; and (2) the lack of common invasibility metrics, as illustrated by our review of all invasibility-related papers published in 2013. To facilitate both cross-habitat comparison and more robust ecological generalizations, we clarify the definitions of invasibility and DI, and for the first time propose a common metric for quantifying invasibility based on a habitat's resource availability as inferred from relative resident species richness and biomass. We demonstrate the feasibility of our metric using empirical data collected from 2475 plots from three forest ecosystems in the eastern United States. We also propose a similar metric for DI. Our unified, resource-based metrics are scaled from 0 to 1, facilitating cross-habitat comparisons. Our proposed metrics clearly distinguish invasibility and DI from each other, which will help to (1) advance invasion ecology by allowing more robust testing of generalizations and (2) facilitate more effective invasive species control and management.