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A systematic review of the effects of climate variability and change on black and brown bear ecology and interactions with humans

Climate change poses a pervasive threat to humans and wildlife by altering resource availability, changing co-occurrences, and directly or indirectly influencing human-wildlife interactions. For many wildlife agencies in North America, managing bears ( Ursus spp.) and human-bear interactions is a priority, yet the direct and indirect effects of climate change are exacerbating management challenges. Understanding the underlying ecological drivers of bear responses to climate variability and change, and the implications for conflict, will be critical for maintaining human-bear coexistence in North America. We synthesized 120 articles that identified direct and indirect mechanisms by which climate variability and change affect brown bears ( Ursus arctos ) and American black bears ( Ursus americanus ) in North America. The literature focused on examining climate impacts on bear diet, body size, habitat selection, space use, activity, denning chronology, and population demographics and dynamics. Across these categories, we summarized the documented and projected bear responses and resulting implications for human-bear interactions. Climate-driven changes in natural food availability were frequently implicated in influencing bear behavior and demography, and creating conditions under which interactions with humans are likely to increase. Bears in North America may face increased challenges as habitat and natural food availability continue to be altered by climate change. Our review provides a foundation upon which to identify climate drivers of bear ecology, conditions conducive to human-bear interactions, and adaptive management strategies. Given substantial evidence of climate impacts to bears, incorporating climate considerations into bear management can help managers strategically allocate resources and promote human-bear coexistence.

Biological Conservation↗

Improving species status assessments under the U.S. Endangered Species Act and implications for multispecies conservation challenges worldwide

Despite its successes, the U.S. Endangered Species Act (ESA) has proven challenging to implement due to funding limitations, workload backlog, and other problems. As threats to species survival intensify and as more species come under threat, the need for the ESA and similar conservation laws and policies in other countries to function efficiently has grown. Attempts by the U.S. Fish and Wildlife Service (USFWS) to streamline ESA decisions include multispecies recovery plans and habitat conservation plans. We address species status assessment (SSA), a USFWS process to inform ESA decisions from listing to recovery, within the context of multispecies and ecosystem planning. Although existing SSAs have a single-species focus, ecosystem-based research can efficiently inform multiple SSAs within a region and provide a foundation for transition to multispecies SSAs in the future. We considered at-risk grassland species and ecosystems within the southeastern United States, where a disproportionate number of rare and endemic species are associated with grasslands. To initiate our ecosystem-based approach, we used a combined literature-based and structured World Café workshop format to identify science needs for SSAs. Discussions concentrated on 5 categories of threats to grassland species and ecosystems, consistent with recommendations to make shared threats a focus of planning under the ESA: (1) habitat loss, fragmentation, and disruption of functional connectivity; (2) climate change; (3) altered disturbance regimes; (4) invasive species; and (5) localized impacts. For each threat, workshop participants identified science and information needs, including database availability, research priorities, and modeling and mapping needs. Grouping species by habitat and shared threats can make the SSA process and other planning processes for conservation of at-risk species worldwide more efficient and useful. We found a combination of literature review and structured discussion effective for identifying the scientific information and analysis needed to support the development of multiple SSAs.

Conservation Biology↗

Africa’s drylands in a changing world: Challenges for wildlife conservation under climate and land-use changes in the Greater Etosha Landscape

Proclaimed in 1907, Etosha National Park in northern Namibia is an iconic dryland system with a rich history of wildlife conservation and research. A recent research symposium on wildlife conservation in the Greater Etosha Landscape (GEL) highlighted increased concern of how intensification of global change will affect wildlife conservation based on participant responses to a questionnaire. The GEL includes Etosha and surrounding areas, the latter divided by a veterinary fence into large, private farms to the south and communal areas of residential and farming land to the north. Here, we leverage our knowledge of this ecosystem to provide insight into the broader challenges facing wildlife conservation in this vulnerable dryland environment. We first look backward, summarizing the history of wildlife conservation and research trends in the GEL based on a literature review, providing a broad-scale understanding of the socioecological processes that drive dryland system dynamics. We then look forward, focusing on eight key areas of challenge and opportunity for this ecosystem: climate change, water availability and quality, vegetation and fire management, adaptability of wildlife populations, disease risk, human-wildlife conflict, wildlife crime, and human dimensions of wildlife conservation. Using this model system, we summarize key lessons and identify critical threats highlighting future research needs to support wildlife management. Research in the GEL has followed a trajectory seen elsewhere reflecting an increase in complexity and integration across biological scales over time. Yet, despite these trends, a gap exists between the scope of recent research efforts and the needs of wildlife conservation to adapt to climate and land-use changes. Given the complex nature of climate change, in addition to locally existing system stressors, a framework of forward-thinking adaptive management to address these challenges, supported by integrative and multidisciplinary research could be beneficial. One critical area for growth is to better integrate research and wildlife management across land-use types. Such efforts have the potential to support wildlife conservation efforts and human development goals, while building resilience against the impacts of climate change. While our conclusions reflect the specifics of the GEL ecosystem, they have direct relevance for other African dryland systems impacted by global change.

Africa, Greater Etosha Landscape↗

Surveys for presence of Oregon spotted frog (Rana pretiosa): background information and field methods

The Oregon spotted frog (Rana pretiosa) is the most aquatic of the native frogs in the Pacific Northwest. The common name derives from the pattern of black, ragged-edged spots set against a brown or red ground color on the dorsum of adult frogs. Oregon spotted frogs are generally associated with wetland complexes that have several aquatic habitat types and sizeable coverage of emergent vegetation. Like other ranid frogs native to the Northwest, Oregon spotted frogs breed in spring, larvae transform in summer of their breeding year, and adults tend to be relatively short lived (3-5 yrs). Each life stage (egg, tadpole, juvenile and adult) has characteristics that present challenges for detection. Breeding can be explosive and completed within 1-2 weeks. Egg masses are laid in aggregations, often in a few locations in large areas of potential habitat. Egg masses can develop, hatch, and disintegrate in <2 weeks during warm weather. Tadpoles can be difficult to identify, have low survival, and spend most of their 3-4 months hidden in vegetation or flocculant substrates. Juveniles and adults are often difficult to capture and can spend summers away from breeding areas. Moreover, a substantial portion of extant populations are of limited size (<100 breeding adults), and field densities of all life stages are often low. An understanding of the biology of the species and use of multiple visits are thus important for assessing presence of Oregon spotted frogs. This report is meant to be a resource for USDA Region 6 Forest Service (FS) and OR/WA Bureau of Land Management (BLM) personnel tasked with surveying for the presence of Oregon spotted frogs. Our objective was to summarize information to improve the efficiency of field surveys and increase chances of detection if frogs are present. We include overviews of historical and extant ranges of Oregon spotted frog. We briefly summarize what is known of Oregon spotted frog habitat associations and review aspects of behavior and ecology that are likely to affect detectability in the field. We summarize characteristics that can help differentiate Oregon spotted frog life stages from other northwestern ranid frogs encountered during surveys. Appendices include examples of data collection formats and a protocol for disinfecting field gear.

Interagency Special Status / Sensitive Species Pro↗

Global positioning system and associated technologies in animal behaviour and ecological research

Biologists can equip animals with global positioning system (GPS) technology to obtain accurate (less than or equal to 30 m) locations that can be combined with sensor data to study animal behaviour and ecology. We provide the background of GPS techniques that have been used to gather data for wildlife studies. We review how GPS has been integrated into functional systems with data storage, data transfer, power supplies, packaging and sensor technologies to collect temperature, activity, proximity and mortality data from terrestrial species and birds. GPS 'rapid fixing' technologies combined with sensors provide location, dive frequency and duration profiles, and underwater acoustic information for the study of marine species. We examine how these rapid fixing technologies may be applied to terrestrial and avian applications. We discuss positional data quality and the capability for high-frequency sampling associated with GPS locations. We present alternatives for storing and retrieving data by using dataloggers (biologging), radio-frequency download systems (e.g. very high frequency, spread spectrum), integration of GPS with other satellite systems (e.g. Argos, Globalstar) and potential new data recovery technologies (e.g. network nodes). GPS is one component among many rapidly evolving technologies. Therefore, we recommend that users and suppliers interact to ensure the availability of appropriate equipment to meet animal research objectives.

Philosophical Transactions of the Royal Society B:↗

Lead in the environment

Anthropogenic uses of lead have probably altered its availability and environmental distribution more than any other toxic element. Consequently, lead concentrations in many living organisms may be approaching thresholds of toxicity for the adverse effects of lead. Such thresholds are difficult to define, as they vary with the chemical and physical form of lead, exposure regime, other elements present and also vary both within and between species. The technological capability to accurately quantify low lead concentrations has increased over the last decade, and physiological and behavioral effects have been measured in wildlife with tissue lead concentrations below those previously considered safe for humans.s.236 Consequently. lead criteria for the protection of wildlife and human health are frequently under review, and 'thresholds' of lead toxicity are being reconsidered. Proposed lead criteria for the protection of natural resources have been reviewed by Eisler. Uptake of lead by plants is limited by its generally low availability in soils and sediments, and toxicity may be limited by storage mechanisms and its apparently limited translocation within most plants. Lead does not generally accumulate within the foliar parts of plants, which limits its transfer to higher trophic levels. Although lead may concentrate in plant and animal tissues, no evidence of biomagnification exists. Acid deposition onto surface waters and soils with low buffering capacity may influence the availability of lead for uptake by plants and animals, and this may merit investigation at susceptible sites. The biological significance of chronic low-level lead exposure to wildlife is sometimes difficult to quantify. Animals living in urban environments or near point sources of lead emission are inevitably subject to greater exposure to lead and enhanced risk of lead poisoning. Increasingly strict controls on lead emissions in many countries have reduced exposure to lead from some sources. and the .reduction of lead in gasoline has resulted in lower tissue lead concentrations in humans and wildlife from many, particularly urban, locations. However, it has been suggested that increasing use of organic lead compounds as catalysts for the production of plastics and as wood preservatives and biocides could adversely affect wildlife. The most significant source of direct wildlife mortality from lead is spent gunshot and fishing sinkers. Elevated mortality from shot ingestion in avian species resulted in the introduction of nontoxic (steel) shot zones along certain flyways in the United States in the mid-1970s and a total ban on the use of lead for waterfowl and coot hunting nationwide by 1992. Several other countries are now following suit and have either banned or are in the process of restricting the use of lead shot for waterfowl hunting. In the United States it has been estimated that since the 1986 hunting season. when the use of nontoxic shot became widespread. over 6 million ducks have not been lost to lead poisoning. Raptors, especially eagles, have also apparently benefited. although lead poisoning from ingestion of bullet fragments remains a problem for the critically threatened California condor. Quantifying reductions in lead mortality rates would be difficult since eagle populations throughout North America are rapidly recovering from other anthropogenic perturbations, especially organochlorine pesticides.

Book chapter↗

Assessing the ecological risks of per‐ and polyfluoroalkyl substances: Current state‐of‐the science and a proposed path forward

Per‐ and poly‐fluoroalkyl substances (PFAS) encompass a large, heterogenous group of chemicals of potential concern to human health and the environment. Based on information for a few relatively well‐understood PFAS such as perfluorooctane sulfonate and perfluorooctanoate, there is ample basis to suspect that at least a subset can be considered persistent, bioaccumulative, and/or toxic. However, data suitable for determining risks in either prospective or retrospective assessments are lacking for the majority of PFAS. In August 2019, the Society of Environmental Toxicology and Chemistry sponsored a workshop that focused on the state‐of‐the‐science supporting risk assessment of PFAS. The present review summarizes discussions concerning the ecotoxicology and ecological risks of PFAS. First, we summarize currently available information relevant to problem formulation/prioritization, exposure, and hazard/effects of PFAS in the context of regulatory and ecological risk assessment activities from around the world. We then describe critical gaps and uncertainties relative to ecological risk assessments for PFAS and propose approaches to address these needs. Recommendations include the development of more comprehensive monitoring programs to support exposure assessment, an emphasis on research to support the formulation of predictive models for bioaccumulation, and the development of in silico, in vitro, and in vivo methods to efficiently assess biological effects for potentially sensitive species/endpoints. Addressing needs associated with assessing the ecological risk of PFAS will require cross‐disciplinary approaches that employ both conventional and new methods in an integrated, resource‐effective manner.

Environmental Toxicology and Chemistry↗

What processes must we understand to forecast regional-scale population dynamics?

An urgent challenge facing biologists is predicting the regional-scale population dynamics of species facing environmental change. Biologists suggest that we must move beyond predictions based on phenomenological models and instead base predictions on underlying processes. For example, population biologists, evolutionary biologists, community ecologists and ecophysiologists all argue that the respective processes they study are essential. Must our models include processes from all of these fields? We argue that answering this critical question is ultimately an empirical exercise requiring a substantial amount of data that have not been integrated for any system to date. To motivate and facilitate the necessary data collection and integration, we first review the potential importance of each mechanism for skilful prediction. We then develop a conceptual framework based on reaction norms, and propose a hierarchical Bayesian statistical framework to integrate processes affecting reaction norms at different scales. The ambitious research programme we advocate is rapidly becoming feasible due to novel collaborations, datasets and analytical tools.

Proceedings of the Royal Society B: Biological Sci↗

Streamside habitats in southern forested wetlands: Their role and implications for management

The value of streamside forests to fish and wildlife and the influence of forest management on their value have been recognized in a general sense for decades. However, in today’s climate of increasing environmental regulation and intensive forest management, there is need for more detailed understanding of the value of streamside forests to fish and wildlife. Dickson and Huntley (1987:38) described the problem well when they wrote that “quantitative data on the effects of riparian zones on wildlife populations are insufficient to enable wildlife managers to justify the retention of riparian zones in land-use plans on a biological and economical basis.” Due in large part to the passage of water pollution control legislation, as well as legislation mandating multiple-use management in our national forests, progress is being made. During the last l&15 years, a great deal of research has been directed at understanding the value and appropriate management of the riparian zone (Brouha and Parsons 1985). The vast majority of research on riparian habitats has been conducted in western forests and/or narrow zones in otherwise upland areas. Also, much of the work done in southern forested wetlands has applied to entire floodplain forests. In this paper, we review the literature on streamside habitats within southern forested wetlands and, for reasons described in the next section, we make a distinction between streamside forests and floodplain (or riparian) forests. We also discuss in less detail the value of streamside habitats within other southern forest types, such as pine or mixed pine-hardwood.

General Technical Report↗

Reptiles and amphibians

Summary – We reviewed all the peer-reviewed scientific publications we could find on the known and potential effects of wind farm development, operation, maintenance, and decommissioning on reptiles and amphibians (collectively herpetofauna) worldwide. Both groups are declining globally due to a multitude of threats including energy development. Effect studies were limited to the long-term research by the authors on Agassiz’s Desert Tortoise ecology and behavior at single operational wind farm in California, US and an analysis of the effects of wind farm installation on species richness of vertebrates including reptiles and amphibians in northwestern Portugal. Research on Agassiz’s Desert Tortoise found few demonstrable differences in biological parameters between populations in the wind farm and those in more natural habitats. High reproductive output is due to the regional climate and not to the presence or operation of the wind farm. Site operations have resulted in death and injury to a small number of adult tortoises and over the long-term tortoises now appear to avoid the areas of greatest turbine concentration. Research in Portugal using models and simulations based on empirical data show that vertebrate species richness (including herpetofauna) decreased by almost 20% after the installation of only two large monopole turbines per 250 x 250 m plot. Knowledge of the known responses of herpetofauna to various disturbances allows identification of potential impacts from construction material acquisition in offsite areas, mortality and stress due to impacts of roads and related infrastructure, destruction and modification of habitat, habitat fragmentation and barriers to gene flow, noise, vibration, electromagnetic field generation, heat from buried high voltage transmission lines, alteration of local and regional climate, predator attraction, and increased risk of fire. Research on herpetofauna lags far behind what is needed and, in particular, before-after-control-impact studies are critically needed to identify cause and effect relationships in order to develop effective mitigation strategies for any negative impacts.

Book chapter↗

Conservation implications when the nest predators are known

Conservation and management of passerines has largely focused on habitat manipulation or restoration because the natural communities on which these birds depend have been destroyed and fragmented. However, productivity is another important aspect of avian conservation, and nest predation can be a large source of nesting mortality for passerines. Recent studies using video surveillance to identify nest predators allow researchers to start evaluating what methods could be used to mitigate nest predation to help passerines of conservation concern. From recent studies, we identified latitudinal and habitat-related patterns in the importance of predator groups that depredate passerine nests. We then reviewed how knowledge of specific nest predators can benefit conservation of bird species of concern. Mammals were the dominant predator group in northern grasslands. Snakes were the dominant predator group in southern habitats. Fire ants were only a nest predator in southern latitudes. Differences in the importance of predator species or groups were likely the result of both their geographic patterns of distribution and habitat preferences. Some direct and indirect predator control measures developed for waterfowl management potentially could be used to benefit passerine productivity. We reviewed three examples-cowbirds, snakes in shrublands, and ground squirrels in grasslands-to illustrate how different predator control strategies may be needed in different situations. Mitigation of passerine nest predation will need to be based on knowledge of predator communities to be effective. This requires large samples of predation events with identified predators; video technology is essential for this task.

Studies in Avian Biology↗

Implementation of a framework for multi-species, multi-objective adaptive management in Delaware Bay

Decision analytic approaches have been widely recommended as well suited to solving disputed and ecologically complex natural resource management problems with multiple objectives and high uncertainty. However, the difference between theory and practice is substantial, as there are very few actual resource management programs that represent formal applications of decision analysis. We applied the process of structured decision making to Atlantic horseshoe crab harvest decisions in the Delaware Bay region to develop a multispecies adaptive management (AM) plan, which is currently being implemented. Horseshoe crab harvest has been a controversial management issue since the late 1990s. A largely unregulated horseshoe crab harvest caused a decline in crab spawning abundance. That decline coincided with a major decline in migratory shorebird populations that consume horseshoe crab eggs on the sandy beaches of Delaware Bay during spring migration. Our approach incorporated multiple stakeholders, including fishery and shorebird conservation advocates, to account for diverse management objectives and varied opinions on ecosystem function. Through consensus building, we devised an objective statement and quantitative objective function to evaluate alternative crab harvest policies. We developed a set of competing ecological models accounting for the leading hypotheses on the interaction between shorebirds and horseshoe crabs. The models were initially weighted based on stakeholder confidence in these hypotheses, but weights will be adjusted based on monitoring and Bayesian model weight updating. These models were used together to predict the effects of management actions on the crab and shorebird populations. Finally, we used a dynamic optimization routine to identify the state dependent optimal harvest policy for horseshoe crabs, given the possible actions, the stated objectives and our competing hypotheses about system function. The AM plan was reviewed, accepted and implemented by the Atlantic States Marine Fisheries Commission in 2012 and 2013. While disagreements among stakeholders persist, structured decision making enabled unprecedented progress towards a transparent and consensus driven management plan for crabs and shorebirds in Delaware Bay.

Biological Conservation↗

Climate-smart invasive species management for 21st century global change challenges

Addressing the global challenges of climate change and biotic invasions requires understanding their interactions and implications for natural resource management. To facilitate and support invasive species management in a changing climate, we review how climate change and invasions interact to impact the planning, action, and outcomes of invasive species management. Climate change is affecting which species natural resource managers need to assess, monitor, and target by facilitating the introduction of new potential invasive species and altering pathways of introduction and spread. Climate-driven shifts in seasonal conditions and phenological events disrupt and require more flexible management timelines. Climate change may reduce the efficacy and feasibility of current treatment methods and create a need to shift management goals as native ecosystems become more vulnerable to disturbance. Additionally, extreme climate events can compound the spread and impacts of biological invasions, making invasive species management a necessary part of extreme event preparation and response planning. As a solution to these challenges, we propose climate-smart invasive species management, which we define as the approaches that managers and decision-makers can take to address the interactive effects of climate change and invasions. Climate-smart invasive species management includes considering shifts in species ranges, abundance, and impacts to inform monitoring, treatment, and regulation of potential and new invaders at multiple scales. Managers can also adjust the timing of and approach to treatment as efficacy and feasibility change, promote resilient ecosystems through climate-smart restoration, and consider the effects of climate change when setting management goals. Furthermore, climate adaptation funds could expand avenues of funding and support for invasive species management. Explicitly considering the interactions of climate change and biological invasions within organizational decision-making and policy will lead to more effective management and can promote more resilient landscapes.

Global Change Biology↗

Type specimens of Crotalus scutulatus (Chordata: Reptilia: Squamata: Viperidae) re-examined, with new evidence after more than a century of confusion

The original description of Crotalus scutulatus (Chordata: Reptilia: Squamata: Viperidae) was published in 1861 by Robert Kennicott, who did not identify a type specimen or a type locality. We review the history of specimens purported to be the type(s) and various designations of type locality. We provide evidence that ANSP 7069 (formerly one of two specimens of USNM 5027) is the holotype and that the appropriate type locality is Fort Buchanan, near present-day Sonoita, in Santa Cruz County, Arizona.

Proceedings of the Biological Society of Washingto↗

Characterization of the water resources of the Pamunkey River watershed in Virginia—A review of water science, management, and traditional ecological knowledge

In central Virginia, the Pamunkey Indian Tribe and Reservation are facing increasingly complex water resource issues related to quantity and quality. Documentation of surface-water, groundwater, water quality, land subsidence, sea-level rise, and river ecology issues in the Pamunkey River watershed and incorporation of traditional ecological knowledge into these research topics may improve understanding of the water resources broadly. This report summarizes the relevant traditional ecological knowledge and scientific literature to elucidate gaps in the total combined knowledge of a suite of water science topics concerning the Pamunkey River watershed. This suite of water science topics includes some of the issues that the Pamunkey Indian Tribe and Reservation are facing within the watershed: fragmentation of streams and management of streamflow, flooding, degrading water quality, groundwater extraction, relative sea-level rise, rapid changes in ecosystems, loss of ecological and biological diversity, and climate change. Gaps in total combined knowledge are pervasive throughout each of these water resource topics. Identifying these knowledge gaps can help inform future research and management strategies.

Virginia↗

Sound and sturgeon: Bioacoustics and anthropogenic sound

Sturgeons are basal bony fishes, most species of which are considered threatened and/or endangered. Like all fishes, sturgeons use hearing to learn about their environment and perhaps communicate with conspecifics, as in mating. Thus, anything that impacts the ability of sturgeon to hear biologically important sounds could impact fitness and survival of individuals and populations. There is growing concern that the sounds produced by human activities (anthropogenic sound), such as from shipping, commercial barge navigation on rivers, offshore windfarms, and oil and gas exploration, could impact hearing by aquatic organisms. Thus, it is critical to understand how sturgeon hear, what they hear, and how they use sound. Such data are needed to set regulatory criteria for anthropogenic sound to protect these animals. However, very little is known about sturgeon behavioral responses to sound and their use of sound. To help understand the issues related to sturgeon and anthropogenic sound, this review first examines what is known about sturgeon bioacoustics. It then considers the potential effects of anthropogenic sound on sturgeon and, finally identifies areas of research that could substantially improve knowledge of sturgeon bioacoustics and effects of anthropogenic sound. Filling these gaps will help regulators establish appropriate protection for sturgeon.

The Journal of the Acoustical Society of America↗

Wing pathology of white-nose syndrome in bats suggests life-threatening disruption of physiology

White-nose syndrome (WNS) is causing unprecedented declines in several species of North American bats. The characteristic lesions of WNS are caused by the fungus Geomyces destructans , which erodes and replaces the living skin of bats while they hibernate. It is unknown how this infection kills the bats. We review here the unique physiological importance of wings to hibernating bats in relation to the damage caused by G. destructans and propose that mortality is caused by catastrophic disruption of wing-dependent physiological functions. Mechanisms of disease associated with G. destructans seem specific to hibernating bats and are most analogous to disease caused by chytrid fungus in amphibians.

BMC Biology↗

Exploring the role of cryptic nitrogen fixers in terrestrial ecosystems: A frontier in nitrogen cycling research

Biological nitrogen fixation represents the largest natural flux of new nitrogen (N) into terrestrial ecosystems, providing a critical N source to support net primary productivity of both natural and agricultural systems. When they are common, symbiotic associations between plants and bacteria can add more than 100 kg N ha −1 y −1 to ecosystems. Yet, these associations are uncommon in many terrestrial ecosystems. In most cases, N inputs derive from more cryptic sources, including mutualistic and/or free-living microorganisms in soil, plant litter, decomposing roots and wood, lichens, insects, and mosses, among others. Unfortunately, large gaps remain in the understanding of cryptic N fixation. We conducted a literature review to explore rates, patterns, and controls of cryptic N fixation in both unmanaged and agricultural ecosystems. Our analysis indicates that, as is common with N fixation, rates are highly variable across most cryptic niches, with N inputs in any particular cryptic niche ranging from near zero to more than 20 kg ha −1 y −1 . Such large variation underscores the need for more comprehensive measurements of N fixation by organisms not in symbiotic relationships with vascular plants in terrestrial ecosystems, as well as identifying the factors that govern cryptic N fixation rates. We highlight several challenges, opportunities, and priorities in this important research area, and we propose a conceptual model that posits an interacting hierarchy of biophysical and biogeochemical controls over N fixation that should generate valuable new hypotheses and research.

Ecosystems↗