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Assessing the components of adaptive capacity to improve conservation and management efforts under global change

Natural-resource managers and other conservation practitioners are under unprecedented pressure to categorize and quantify the vulnerability of natural systems based on assessment of the exposure, sensitivity, and adaptive capacity of species to climate change. Despite the urgent need for these assessments, neither the theoretical basis of adaptive capacity nor the practical issues underlying its quantification has been articulated in a manner that is directly applicable to natural-resource management. Both are critical for researchers, managers, and other conservation practitioners to develop reliable strategies for assessing adaptive capacity. Drawing from principles of classical and contemporary research and examples from terrestrial, marine, plant, and animal systems, we examined broadly the theory behind the concept of adaptive capacity. We then considered how interdisciplinary, trait- and triage-based approaches encompassing the oft-overlooked interactions among components of adaptive capacity can be used to identify species and populations likely to have higher (or lower) adaptive capacity. We identified the challenges and value of such endeavors and argue for a concerted interdisciplinary research approach that combines ecology, ecological genetics, and eco-physiology to reflect the interacting components of adaptive capacity. We aimed to provide a basis for constructive discussion between natural-resource managers and researchers, discussions urgently needed to identify research directions that will deliver answers to real-world questions facing resource managers, other conservation practitioners, and policy makers. Directing research to both seek general patterns and identify ways to facilitate adaptive capacity of key species and populations within species, will enable conservation ecologists and resource managers to maximize returns on research and management investment and arrive at novel and dynamic management and policy decisions.

Conservation Biology

Exposure and effects of bioaccumulative contaminants of emerging concern in tree swallows nesting across the Laurentian Great Lakes

Contaminants of emerging concern (CECs) are a loosely defined group of chemicals whose wide-spread usage or presence in the environment has occurred more recently or for which there has been relatively little research done until recently. Many of these CECs are not currently regulated. The National Toxicology Program within the U.S. Department of Health and Human Services estimates that about 2000 CECs are introduced each year (https://ntp.niehs.nih.gov/about/). An unknown number may pose a risk to human or animal health. The Phase 1 (2010 – 2014) CEC work in birds, which is the subject of this report, assessed exposure across the Great Lakes to polybrominated diphenyl ethers (PBDEs), perfluorinated compounds (PFASs), and polycyclic aromatic hydrocarbons (PAHs), and put those exposures into context with data from biologically relevant endpoints such as reproductive success, as well as, physiological response indicators (bioindicators) to assess possible effects. The group of chemicals included in Phase 1 were mainly those chemicals that bioaccumulate in tissues. Phase 2 (2015 – 2019) CEC work with tree swallows was expanded to include CECs whose occurrence in the environment is more temporary or seasonal, and that do not necessarily bioaccumulate. These are often called pseudo-persistent, because, while they are not long-lived in the environment, there are often daily inputs via waste water treatment plants, and run-off from farm fields and storm drainages, thereby making them available to biota year-round. These include pharmaceuticals, personal care products, and newer pesticides including herbicides. Tree swallow work on these less persistent CECs will be reported in the future, however see other Appendices in this report for information on some of these types of CECs (Appendices A, B, D).

Laurentian Great Lakes

Environmental gradients of selection for an alpine-obligate bird, the white-tailed ptarmigan (Lagopus leucura)

The warming climate will expose alpine species adapted to a highly seasonal, harsh environment to novel environmental conditions. A species can shift their distribution, acclimate, or adapt in response to a new climate. Alpine species have little suitable habitat to shift their distribution, and the limits of acclimation will likely be tested by climate change in the long-term. Adaptive genetic variation may provide the raw material for species to adapt to this changing environment. Here, we use a genomic approach to describe adaptive divergence in an alpine-obligate species, the white-tailed ptarmigan ( Lagopus leucura ), a species distributed from Alaska to New Mexico, across an environmentally variable geographic range. Previous work has identified genetic structure and morphological, behavioral, and physiological differences across the species’ range; however, those studies were unable to determine the degree to which adaptive divergence is correlated with local variation in environmental conditions. We used a genome-wide dataset generated from 95 white-tailed ptarmigan distributed throughout the species’ range and genotype–environment association analyses to identify the genetic signature and environmental drivers of local adaptation. We detected associations between multiple environmental gradients and candidate adaptive loci, suggesting ptarmigan populations may be locally adapted to the plant community composition, elevation, local climate, and to the seasonality of the environment. Overall, our results suggest there may be groups within the species’ range with genetic variation that could be essential for adapting to a changing climate and helpful in guiding conservation action.

Alaska, Washington, Montana, Colorado, New Mexico,

Multi-year cut-to-drown management limits Phragmites australis growth, belowground resources, and rhizome viability in Great Lakes wetlands

The distribution and abundance of Phragmites in the Great Lakes coastal zone has expanded in part due to its unique ventilation physiology and its ability to take advantage of changes in lake levels over the past several decades. During an extended period of low lake levels in the early 2000s, Phragmites expanded into vast shallow water areas as lake bottoms were exposed. Many of those populations were able to persist when lake levels rose several years later. Here, we investigate a management strategy, known as cut-to-drown, that takes advantage of elevated water levels to effectively control Phragmites by cutting stems underwater, thereby drowning the plant and depleting its stored resources. We tested the impacts of cut timing and frequency on the effectiveness of the cut-to-drown strategy through a multi-year manipulative field study. After two seasons of treatments, we found a 92–99 % reduction in stem density, depending on cut timing. Carbohydrate reserves in rhizomes declined 75–92 %, such that rhizome viability was reduced by 73–100 %, depending on cut timing. We found that cutting continuously (every 2–3 weeks) throughout two growing seasons was the most effective way to implement cut-to-drown but provide evidence that less frequent cutting could produce similar results. Our study provides important evidence to inform best management practices for cut-to-drown. Particularly as lake level fluctuations become more frequent and extreme in the future, cut-to-drown could serve as a useful tool for limiting Phragmites’ ability to persist in high water environments.

Michigan, Ohio

Characterization of ecological risks at the Milltown Reservoir-Clark Fork River Sediments Superfund Site, Montana

A comprehensive field and laboratory approach to the ecological risk assessment for the Milltown Reservoir-Clark Fork River Sediments Site, a Superfund site in the Rocky Mountains of Montana, has been described in the preceding reports of this series. The risk assessment addresses concerns over the ecological impacts of upstream releases of mining wastes to fisheries of the upper Clark Fork River (CFR) and the benthic and terrestrial habitats further downstream in Milltown Reservoir. The risk characterization component of the process integrated results from a triad of information sources: (a) chemistry studies of environmental media to identify and quantify exposures of terrestrial and aquatic organisms to site-related contaminants; (b) ecological or population studies of terrestrial vegetation, birds, benthic communities, and fish; and (c) in situ and laboratory toxicity studies with terrestrial and aquatic invertebrates and plants, small mammals, amphibians, and fish exposed to contaminated surface water, sediments, wetland soils, and food sources. Trophic transfer studies were performed on waterfowl, mammals, and predatory birds using field measurement data on metals concentrations in environmental media and lower trophic food sources. Studies with sediment exposures were incorporated into the Sediment Quality Triad approach to evaluate risks to benthic ecology. Overall results of the wetland and terrestrial studies suggested that acute adverse biological effects were largely absent from the wetland; however, adverse effects to reproductive, growth, and physiological end points of various terrestrial and aquatic species were related to metals exposures in more highly contaminated depositional areas. Feeding studies with contaminated diet collected from the upper CFR indicated that trout are at high risk from elevated metals concentrations in surface water, sediment, and aquatic invertebrates. Integration of chemical analyses with toxicological and ecological evaluations of metal effects on the wetland and fishery has provided an important foundation for environmental decisions at this site.

Montana

Endothal derivatives as aquatic herbicides in fishery habitats

The disodium salt of 3,6-endoxohexahydrophthalic acid (disodium endothal) and the derivative identified by the manufacturer as the di- N,N ′-dimethylococoamine salt of endothal (coded as TD-47) were particularly effective upon submersed species of aquatic vegetation as contact herbicides. Disodium endothal at concentrations of 0.5 to 10.0 ppmw was effective in controlling approximately 50 per cent of the 19 species of plants involved in 270 tests. TD-47 at concentrations of 0.02 to 10.0 ppmw trolled 77 per cent of the 11 plant species in 94 tests. Algae ( Chara, Cladophora, Pithophora , and Spirogyra ) were more effectively controlled by TD-47 than by disodium endothal. Although TD-47 was at least 10 times more herbicidal than disodium endothal, it was about 100 times more toxic to fish. Disodium endothal was more than 50 per cent effective on submersed aquatic plants at rates in excess of 2.5 ppmw with a wide margin of safety in fish (4- to 10-fold). Disodium endothal had a median tolerance limit ranging from 95 to 150 ppmw in the aggregate of nine fish species tested extensively. Median tolerance limits for TD-47 ranged from about 0.06 to 0.3 ppmw for five species of fish. TD-47 applied at a concentration lethal to fish (0.3 to 1.0 ppmw) was effective as a dual management tool in controlling vegetation and achieving partial or complete renovation of stunted fish populations. Young, growing vegetation was most susceptible to control, and best results were achieved at water temperatures exceeding 60 F. Higher rates were required to kill plants as they matured and stands became dense. Endothal liquid formulations were superior to granules in controlling algal mats, floating and emergent plants. Granules were more effective on submersed rooted plants. TD-47 residues were of short duration. The rate of disappearance depended on time and concentration. Detectable residues disappeared within 8 days following application of 0.3 ppmw and within 2 weeks for 0.6 ppmw. However, 1.0 to 3.0 ppmw took up to 25 days to disappear. Some residues were found in fish-food organisms from treated enclosures 3 weeks after application. Fish flesh showed no absorption of endothal-armeens at sublethal concentrations. Intraperitoneal injection of endothal into fish produced a disturbance of the osmoregulation. The physiological effect of endothal was measured by chemical analysis of blood serum.

Weeds

Use of "specific" inhibitors in biogeochemistry and microbial ecology

The above statement, although meant to be tongue in cheek, contains an essential truism: all work with inhibitors is inherently suspect. This fact has been known by biochemists for some time. However, use of chemical inhibitors of enzymic systems and membranes continues to be a common approach taken toward unraveling the biochemistry and biophysics of plants, animals, and microorganisms. Various types of “broad-spectrum” biochemical inhibitors (e.g., poisons, respiratory inhibitors, and uncouplers) have been employed by ecologists for many years in order to demonstrate the active participation of microbes in chemical reactions occurring in natural samples (e.g., soils, sediments, and water). In recent years, considerable advances have been made in our understanding of the biochemistry of microorganisms of biogeochemical interest. Concurrent with these advances have been the discoveries of novel types of compounds that will block the metabolism of one particular group of microbes, but have little disruptive effect on other physiological types. Thus, the term “specific inhibitor” has been applied to these types of compounds when they are used to probe the functions of mixed populations of microorganisms. These substances provide powerful experimental tools for investigating the activity and function of certain types of microorganisms in natural samples.

Book chapter

Options for water-level control in developed wetlands

Wetland habitats in the United States currently are lost at a rate of 260,000 acres/year (105,218 ha/year). Consequently, water birds concentrate in fewer and smaller areas. Such concentrations may deplete food supplies and influence behavior, physiology, and survival. Continued losses increase the importance of sound management of the remaining wetlands because water birds depend on them. Human activities modified the natural hydrology of most remaining wetlands in the conterminous United States, and such hydrologic alterations frequently reduce wetland productivity. The restoration of original wetland functions and productivity often requires the development of water distribution and discharge systems to emulate natural hydrologic regimes. Construction of levees and correct placement of control structures and water-delivery and water-discharge systems are necessary to (1) create soil and water conditions for the germination of desirable plants, (2) control nuisance vegetation, (3) promote the production of invertebrates, and (4) make foods available for wildlife that depends of wetlands (Leaflets 13.2.1 and 13.4.6). This paper provides basic guidelines for the design of wetlands that benefit wildlife. If biological considerations are not incorporated into such designs, the capability of managing wetlands for water birds is reduced and costs often are greater. Although we address the development of palustrine wetlands in migration and wintering areas, many of the discussed principles are applicable to the development of other wetland types and in other locations.

Book chapter

Chapter 6: Climate change, wildlife, and wildlife habitats in the Oregon Coast Range

Climate change is likely to have profound effects on wildlife species within the Oregon Coast Adaptation Partnership (OCAP) assessment area, although the direction and magnitude of effects are likely to vary across species. Increased mean and extreme temperatures, especially during summer, may cause shifts in plant and animal species ranges, reduce habitat for some temperature-sensitive wildlife, alter plant phenology and the timing of available food resources, and affect species interactions (e.g., predation, competition). Altered timing of precipitation, summer drought, loss of fog, increased flooding events, earlier snowmelt, and rising sea level may reduce plant productivity, increase tree mortality, shift plant species composition, and lead to reduced wildlife habitat and habitat quality for some forests, riparian areas, wetlands, meadows, estuaries, and beaches. In addition, increasing frequency and extent of wildfire and insect outbreaks may reduce the extent of late-successional forest, reduce habitat connectivity, and increase the spread of invasive species. The biggest change expected for the assessment area is an increase in area where climatic conditions favor coastal mixed forest and a large reduction in area favoring montane conifer forest. Although actual changes in forest types may not necessarily occur by the end of the 21st century, climate change may add physiological and behavioral stress to wildlife. Some wildlife species will be able to persist in place and adapt to new conditions; some may be able to migrate to find suitable habitat; and some may be greatly reduced or extirpated from the assessment area or even go extinct. Shifts in major tree and shrub species will play a major role in food, den, and cover availability for wildlife. Rising sea level will reduce low-elevation habitats along the coast. An increase in the frequency of high-severity weather events will increase frequency and magnitude of flooding, including debris flow events. Coupled with increased temperatures during summer, this may reduce or fragment important ecosystems for aquatic and semi-aquatic species. Evaluation of the vulnerabilities of nine wildlife species based on literature reviews suggests that each species has life-history attributes that can lead to resilience or vulnerability to climate change effects. Depending on long-term objectives, several broad adaptation strategies focus on protecting refugia, establishing redundant wildlife strongholds with large-scale connectivity, and promoting structural and biological complexity.

Oregon

Changes in structure and physiological functioning due to experimentally enhanced precipitation seasonality in a widespread shrub species

Semi-arid shrub steppe occupies a vast geographic range that is characterized in part by distinct seasonal patterns in precipitation. Few studies have evaluated how variability in both the amount and timing of precipitation affect the structure and physiology of shrubs in these systems. We quantified changes in foliar crown parameters, xylem anatomy, gas exchange, and hydraulic transport capacity in deep-rooted Artemisia tridentata shrubs following 20 years of experimental manipulations in amount and seasonal timing of precipitation. We hypothesized that shrub growth (total leaf area per shrub and cover of shrub community), hydraulic transport efficiency, and gas exchange would increase in shrubs in irrigated plots compared to non-irrigated control plots, especially for irrigation applied in winter rather than summer. We also predicted similar changes in xylem anatomy (ring width, vessel size and frequency). Most treatment responses entailed changes in plant structure, and were generally consistent with our hypotheses: total-shrub leaf area, shrub basal area, canopy cover, and maximum sapwood-specific branch hydraulic conductivity were more than 2× greater in shrubs in winter-irrigated compared to control plots, while summer irrigation had few effects on these variables. Irrigation in either season did not affect xylem vessel size, but did increase xylem ring width by ~ 2 × and decreased xylem vessel frequency by about half. Anatomical, morphological, and stand-level abundance of A. tridentata appeared much more responsive to irrigation than state changes in gas exchange, particularly when the extra water is received during winter. Thus, it appears for sagebrush that seasonal timing is at least as important as the amount of precipitation, and that responses to changes in precipitation timing occur through changes in carbon allocation more so than changes in leaf-level carbon gain.

Plant Ecology

Behavior of potentially toxic elements from stoker-boiler fly ash in Interior Alaska: Paired batch leaching and solid-phase characterization

Despite significant investigation of fly ash spills and mineralogical controls on the release of potentially toxic elements (PTEs) from fly ash, interactions with the surficial environment remain relatively poorly understood. We conducted 90-day batch leaching studies with paired analysis of supernatant and solid-phase mineralogy to assess the elemental release and transformation of fly ash upon reaction with aquatic media (18 MΩ cm −1 water and simulated rainwater). The fly ash in this study, collected from the University of Alaska Fairbanks stoker-boiler power plant, is high in unburned carbon (~20% LOI) and highly enriched in several PTEs relative to the upper continental crust. Supernatant concentrations of oxyanion-forming elements (e.g., As, Se, Mo, Sb) remained relatively low and constant, suggesting equilibrium with the solid phase, possibly ettringite [Ca 6 Al 2 (SO 4 ) 3 (OH) 12 •26H 2 O], which is known to incorporate and sorb oxyanion-forming PTEs and was identified by X-ray diffraction. Synthetic precipitation leaching procedure (SPLP) results failed to capture important temporal trends. Lead and Ba supernatant concentrations consistently exceeded drinking water standards, as well as others upon exposure to simulated physiological solutions. Seven-day experiments with dissolved organic matter-isolate solutions indicated that for certain elements, liberation was influenced by carbon concentration and/or the identity of the isolate. Overall, this paired approach can serve as a model for future studies, bridging existing gaps between batch leaching and single-element mineralogical, sorption, or speciation studies.

Alaska

Cadmium toxicity among wildlife in the Colorado Rocky Mountains

Cadmium is known to be both extremely toxic and ubiquitous in natural environments. It occurs in almost all soils, surface waters and plants, and it is readily mobilized by human activities such as mining. As a result, cadmium has been named as a potential health threat to wildlife species; however, because it exists most commonly in the environment as a trace constituent, reported incidences of cadmium toxicity are rare. Here we have measured trace metals in the food web and tissues of white-tailed ptarmigan (Lagopus leucurus) in Colorado. Our results suggest that cadmium toxicity may be more common among natural populations of vertebrates than has been appreciated to date and that cadmium toxicity may often go undetected or unrecognized. In addition, our research shows that ingestion of even trace quantities of cadmium can influence not only the physiology and health of individual organisms, but also the demographics and the distribution of species.

Nature

Ecological acclimation: A framework to integrate fast and slow responses to climate change

Ecological responses to climate change occur across vastly different time-scales, from minutes for physiological plasticity to decades or centuries for community turnover and evolutionary adaptation. Accurately predicting the range of ecosystem trajectories will require models that incorporate both fast processes that may keep pace with climate change and slower ones likely to lag behind and generate disequilibrium dynamics. However, the knowledge necessary for this integration is currently fragmented across disciplines. We develop ‘ecological acclimation’ as a unifying framework to emphasize the similarity of dynamics driven by processes operating on dramatically different time-scales and levels of biological organization. The framework focuses on ecoclimate sensitivities, measured as the change in an ecological response variable per unit of climate change. Acclimation processes acting at different time-scales cause these sensitivities to shift in magnitude and even direction over time. We highlight shifting ecoclimate sensitivities in case studies from diverse ecosystems, including terrestrial plant communities, coral reefs and soil microbiomes. Models predicting future ecosystem states inevitably make assumptions about acclimation processes; these assumptions must be explicit for users to evaluate whether a model is appropriate for a given forecast horizon. Similarly, decision frameworks that clearly account for multiple acclimation processes and their distinct time-scales will help natural resource managers plan for ecological impacts of climate change from years to many decades into the future. We outline a synthetic research programme focused on the time-scales of ecological acclimation to reduce uncertainty in ecological forecasts.

Functional Ecology

Rodentia and lagomorpha

This comprehensive review examines the extensive literature on wild rodents and lagomorphs as biomonitors of environmental contamination. This chapter covers studies dealing with exposure and effects of environmental contaminants on rodent and lagomorph species, including pesticides (organochlorines, organophosphorus and carbamate compounds, herbicides, plant growth regulators, fungicides, and rodenticides), other organic chemicals, metals, radionuclides, and other miscellaneous contaminants. Many research needs become evident when reviewing ecotoxicological data for rodents and lagomorphs, the most striking being the paucity of information on rodent families other than Muridae (mice and rats). While our ability to qualitatively extrapolate effects observed in laboratory studies to field situations is good for a variety of contaminants, quantitative predictions of dose-response relationships are poor because inter-specific variation and differences in exposure patterns between laboratory and wild species to toxicants are for the most part unknown. More sophisticated comparative toxicity studies need to be undertaken that build on previous work in order to develop a database of information, to account for and model differences in exposure pathways, to document interactions among multiple stressors, to generate data establishing thresholds, critical concentrations, and diagnostic guidelines, and even to develop physiologically-based toxicokinetic models. Such efforts may enhance our ability to predict effects on wild populations, including threatened and endangered species.

Book chapter

The physiology of mangrove trees with changing climate

Mangrove forests grow on saline, periodically flooded soils of the tropical and subtropical coasts. The tree species that comprise the mangrove are halophytes that have suites of traits that confer differing levels of tolerance of salinity, aridity, inundation and extremes of temperature. Here we review how climate change and elevated levels of atmospheric CO2 will influence mangrove forests. Tolerance of salinity and inundation in mangroves is associated with the efficient use of water for photosynthetic carbon gain which unpins anticipated gains in productivity with increasing levels of CO2. We review evidence of increases in productivity with increasing CO2, finding that enhancements in growth appear to be similar to trees in non-mangrove habitats and that gains in productivity with elevated CO2 are likely due to changes in biomass allocation. High levels of trait plasticity are observed in some mangrove species, which potentially facilitates their responses to climate change. Trait plasticity is associated with broad tolerance of salinity, aridity, low temperatures and nutrient availability. Because low temperatures and aridity place strong limits on mangrove growth at the edge of their current distribution, increasing temperatures over time and changing rainfall patterns are likely to have an important influence on the distribution of mangroves. We provide a global analysis based on plant traits and IPCC scenarios of changing temperature and aridity that indicates substantial global potential for mangrove expansion.

Book chapter

A combined microbial and ecosystem metric of carbon retention efficiency explains land cover-dependent soil microbial biodiversity–ecosystem function relationships

While soil organic carbon (C) is the foundation of productive and healthy ecosystems, the impact of the ecology of microorganisms on C-cycling remains unknown. We manipulated the diversity, applied here as species richness, of the microbial community present in similar soils on two contrasting land-covers—an adjacent pasture and forest—and observed the transformations of plant detritus and soil organic matter (SOM) using stable isotope ( 13 C) tracing coupled with a novel nuclear magnetic resonance (NMR) experiment. The amount of detritus-C degraded was not affected by the microbial diversity (p > 0.05), however the fate of detritus- and SOM-C across the diversity gradient was complex and land cover-dependent. For example, in the pasture soil, higher diversity led to lower CO 2 production (p = 0.001), a trend driven solely by SOM-C mineralization. There was no relationship between diversity and detritus-C mineralization or production of new mineral-associations after one year (p > 0.05). In contrast, in the forest soil higher diversity resulted in increased detritus-C (p = 0.01) and SOM-C (p = 0.0008) mineralization and decreased mineral-associated organic matter formation (p = 0.02). In both land cover types, retention efficiency—a measure that integrates both microbial physiology and the ability of the ecosystem to retain C—explained C loss and transformation trends. Overall, this demonstrates that the trajectory of C gained and lost is altered by land management-induced changes to microbial communities, soil structure, and chemical characteristics underlying SOM persistence.

Biogeochemistry Letters

Fire and tree death: Understanding and improving modeling of fire-induced tree mortality

Each year wildland fires kill and injure trees on millions of forested hectares globally, affecting plant and animal biodiversity, carbon storage, hydrologic processes, and ecosystem services. The underlying mechanisms of fire-caused tree mortality remain poorly understood, however, limiting the ability to accurately predict mortality and develop robust modeling applications, especially under novel future climates. Virtually all post-fire tree mortality prediction systems are based on the same underlying empirical model described in Ryan and Reinhardt (1988 Can. J. For. Res. 18 1291–7), which was developed from a limited number of species, stretching model assumptions beyond intended limits. We review the current understanding of the mechanisms of fire-induced tree mortality, provide recommended standardized terminology, describe model applications and limitations, and conclude with key knowledge gaps and future directions for research. We suggest a two-pronged approach to future research: (1) continued improvements and evaluations of empirical models to quantify uncertainty and incorporate new regions and species and (2) acceleration of basic, physiological research on the proximate and ultimate causes of fire-induced tree mortality to incorporate processes of tree death into models. Advances in both empirical and process fire-induced tree modeling will allow creation of hybrid models that could advance understanding of how fire injures and kills trees, while improving prediction accuracy of fire-driven feedbacks on ecosystems and landscapes, particularly under novel future conditions.

Environmental Research Letters

Climate change and Arctic ecosystems: 2. Modeling, paleodata-model comparisons, and future projections

Large variations in the composition, structure, and function of Arctic ecosystems are determined by climatic gradients, especially of growing-season warmth, soil moisture, and snow cover. A unified circumpolar classification recognizing five types of tundra was developed. The geographic distributions of vegetation types north of 55°N, including the position of the forest limit and the distributions of the tundra types, could be predicted from climatology using a small set of plant functional types embedded in the biogeochemistry-biogeography model BIOME4. Several palaeoclimate simulations for the last glacial maximum (LGM) and mid-Holocene were used to explore the possibility of simulating past vegetation patterns, which are independently known based on pollen data. The broad outlines of observed changes in vegetation were captured. LGM simulations showed the major reduction of forest, the great extension of graminoid and forb tundra, and the restriction of low- and high-shrub tundra (although not all models produced sufficiently dry conditions to mimic the full observed change). Mid-Holocene simulations reproduced the contrast between northward forest extension in western and central Siberia and stability of the forest limit in Beringia. Projection of the effect of a continued exponential increase in atmospheric CO 2 concentration, based on a transient ocean-atmosphere simulation including sulfate aerosol effects, suggests a potential for larger changes in Arctic ecosystems during the 21st century than have occurred between mid-Holocene and present. Simulated physiological effects of the CO 2 increase (to >700 ppm) at high latitudes were slight compared with the effects of the change in climate.

Journal of Geophysical Research D: Atmospheres