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Rejoinder: Challenge and opportunity in the study of ungulate migration amid environmental change

Increasingly, animals that migrate long distances to exploit seasonal habitats must traverse political boundaries capable of altering the very ecological gradients that promote migratory behavior. This transboundary aspect of migration presents many new challenges and opportunities for research and conservation (e.g., Bolger et al. 2008, Taillon et al. 2012). Work to date has often focused on physical barriers to movement (roads, fences,and housing and energy development) that can threaten migratory populations to varying degrees (Holdo et al. 2011, Sawyer et al. 2013). However, even in the absence of conspicuous barriers, political and jurisdictional boundaries can bring dramatic differences in land use and conservation policy. What happens to migratory populations when these boundaries alter the resources and refuges that they seek on their seasonal journeys?

Ecology

Ecoregions of California

Ecoregions denote areas of general similarity in ecosystems and in the type, quality, and quantity of environmental resources. They are designed to serve as a spatial framework for the research, assessment, management, and monitoring of ecosystems and ecosystem components. By recognizing the spatial differences in the capacities and potentials of ecosystems, ecoregions stratify the environment by its probable response to disturbance (Bryce and others, 1999). These general purpose regions are critical for structuring and implementing ecosystem management strategies across Federal agencies, State agencies, and nongovernment organizations that are responsible for different types of resources in the same geographical areas (Omernik and others, 2000). The approach used to compile this map is based on the premise that ecological regions are hierarchical and can be identified through the analysis of the spatial patterns and the composition of biotic and abiotic phenomena that affect or reflect differences in ecosystem quality and integrity (Wiken, 1986; Omernik, 1987, 1995). These phenomena include geology, physiography, vegetation, climate, soils, land use, wildlife, and hydrology. The relative importance of each characteristic varies from one ecological region to another regardless of the hierarchical level. A Roman numeral hierarchical scheme has been adopted for different levels of ecological regions. Level I is the coarsest level, dividing North America into 15 ecological regions. Level II divides the continent into 50 regions (Commission for Environmental Cooperation Working Group, 1997, map revised 2006). At level III, the continental United States contains 105 ecoregions and the conterminous United States has 85 ecoregions (U.S. Environmental Protection Agency, 2013). Level IV, depicted here for California, is a further refinement of level III ecoregions. Explanations of the methods used to define these ecoregions are given in Omernik (1995), Omernik and others (2000), and Omernik and Griffith (2014). California has great ecological and biological diversity. The State contains offshore islands and coastal lowlands, large alluvial valleys, forested mountain ranges, deserts, and various aquatic habitats. There are 13 level III ecoregions and 177 level IV ecoregions in California and most continue into ecologically similar parts of adjacent States of the United States or Mexico (Bryce and others, 2003; Thorson and others, 2003; Griffith and others, 2014). The California ecoregion map was compiled at a scale of 1:250,000. It revises and subdivides an earlier national ecoregion map that was originally compiled at a smaller scale (Omernik, 1987; U.S. Environmental Protection Agency, 2013). This poster is the result of a collaborative project primarily between U.S. Environmental Protection Agency (USEPA) Region IX, USEPA National Health and Environmental Effects Research Laboratory (Corvallis, Oregon), California Department of Fish and Wildlife (DFW), U.S. Department of Agriculture (USDA)–Natural Resources Conservation Service (NRCS), U.S. Department of the Interior–Geological Survey (USGS), and other State of California agencies and universities. The project is associated with interagency efforts to develop a common framework of ecological regions (McMahon and others, 2001). Reaching that objective requires recognition of the differences in the conceptual approaches and mapping methodologies applied to develop the most common ecoregion-type frameworks, including those developed by the USDA–Forest Service (Bailey and others, 1994; Miles and Goudy, 1997; Cleland and others, 2007), the USEPA (Omernik 1987, 1995), and the NRCS (U.S. Department of Agriculture–Soil Conservation Service, 1981; U.S. Department of Agriculture–Natural Resources Conservation Service, 2006). As each of these frameworks is further refined, their differences are becoming less discernible. Regional collaborative projects such as this one in California, where some agreement has been reached among multiple resource-management agencies, are a step toward attaining consensus and consistency in ecoregion frameworks for the entire nation.

California

The U.S. Geological Survey Flagstaff Science Campus—Providing expertise on planetary science, ecology, water resources, geologic processes, and human interactions with the Earth

T he U.S. Geological Survey’s Flagstaff Science Campus is focused on interdisciplinary study of the Earth and solar system, and has the scientific expertise to detect early environmental changes and provide strategies to minimize possible adverse effects on humanity. The Flagstaff Science Campus (FSC) is located in Flagstaff, Arizona, which is situated in the northern part of the State, home to a wide variety of landscapes and natural resources, including (1) young volcanoes in the San Francisco Volcanic Field, (2) the seven ecological life zones of the San Francisco Peaks, (3) the extensive geologic record of the Colorado Plateau and Grand Canyon, (4) the Colorado River and its perennial, ephemeral, and intermittent tributaries, and (5) a multitude of canyons, mountains, arroyos, and plains. More than 200 scientists, technicians, and support staff provide research, monitoring, and technical advancements in planetary geology and mapping, biology and ecology, Earth-based geology, hydrology, and changing climate and landscapes. Scientists at the FSC work in collaboration with multiple State, Federal, Tribal, municipal, and academic partners to address regional, national, and global environmental issues, and provide scientific outreach to the general public.

Arizona

Decision analysis to advance environmental sustainability

Decision analysis provides a robust framework for complex decisions related to environmental sustainability and conservation, including for energy and water, fisheries and wildlife management, agriculture, and climate change response. The complexities of these problems stem from their large scope and scale, which leads to multiple decision makers, stakeholders, rightsholders, and other entities with potentially competing objectives. These problems often are time limited (e.g., urgent action is required to prevent species’ extinction), involve management interventions over long time scales and delayed responses to management (deep uncertainty), and are impeded by limited resources (funding, capacity, etc.). In this Special Issue on “Decision Analysis to Advance Environmental Sustainability,” we present five case studies of applications of decision analysis to complex problems in environmental sustainability and conservation. These case studies incorporate multiple objectives related to ecological and environmental sustainability, economic and social concerns, and logistics of implementation. They showcase a wide range of tools and applications to these problems. We also provide suggestions for new avenues of research and application of decision analysis to problems of environmental sustainability and conservation, including how to incorporate other decision-making tools into decision analysis processes, how to broaden the reach of decision analysis to other sustainability problems, how to incorporate more stakeholders and rightsholders into the decision process, the potential to incorporate new technology into these processes, identifying more creative alternatives, how to secure more funding, ways to move from decision to action, and how to move beyond status quo to make big transitions necessary to achieve sustainability.

Decision Analysis

Columbia Environmental Research Center

The U.S. Geological Survey Columbia Environmental Research Center performs research to solve challenging environmental problems related to contaminants and habitat alterations in aquatic and terrestrial ecosystems. The research is interdisciplinary and pursued through partnerships within the U.S. Geological Survey and with national, international, state, and local agencies; nongovernmental organizations; and universities. Research is prioritized to provide science to the U.S. Department of the Interior and other natural resource management agencies to inform rehabilitation of degraded habitats and imperiled fish and wildlife populations. The Columbia Environmental Research Center was established in 1966 in Columbia, Missouri, as the U.S. Fish and Wildlife Service’s Fish Pesticide Research Laboratory; the Columbia Environmental Research Center was incorporated into the U.S. Geological Survey in 1996. The U.S. Geological Survey’s staff of 130 includes 90 scientists of which one-half have advanced degrees in ecology, toxicology, biology, biochemistry, chemistry, hydrology, geology, and information technology.

Missouri

The impacts of mangrove range expansion on wetland ecosystem services in the southeastern United States: Current understanding, knowledge gaps, and emerging research needs

Climate change is transforming ecosystems and affecting ecosystem goods and services. Along the Gulf of Mexico and Atlantic coasts of the southeastern United States, the frequency and intensity of extreme freeze events greatly influences whether coastal wetlands are dominated by freeze-sensitive woody plants (mangrove forests) or freeze-tolerant grass-like plants (salt marshes). In response to warming winters, mangroves have been expanding and displacing salt marshes at varying degrees of severity in parts of north Florida, Louisiana, and Texas. As winter warming accelerates, mangrove range expansion is expected to increasingly modify wetland ecosystem structure and function. Because there are differences in the ecological and societal benefits that salt marshes and mangroves provide, coastal environmental managers are challenged to anticipate effects of mangrove expansion on critical wetland ecosystem services, including those related to carbon sequestration, wildlife habitat, storm protection, erosion reduction, water purification, fisheries support, and recreation. Mangrove range expansion may also affect wetland stability in the face of extreme climatic events and rising sea levels. Here, we review current understanding of the effects of mangrove range expansion and displacement of salt marshes on wetland ecosystem services in the southeastern United States. We also identify critical knowledge gaps and emerging research needs regarding the ecological and societal implications of salt marsh displacement by expanding mangrove forests. One consistent theme throughout our review is that there are ecological trade-offs for consideration by coastal managers. Mangrove expansion and marsh displacement can produce beneficial changes in some ecosystem services, while simultaneously producing detrimental changes in other services. Thus, there can be local-scale differences in perceptions of the impacts of mangrove expansion into salt marshes. For very specific local reasons, some individuals may see mangrove expansion as a positive change to be embraced, while others may see mangrove expansion as a negative change to be constrained.

Global Change Biology

Ecotoxicology of wild mammals

An international group of 32 scientists has critically reviewed the scientific literature on exposure and effects of environmental contaminants in wild mammals. Although the absolute number of toxicological studies in domesticated and wild mammals eclipses that for birds, a detailed examination of scientific publications and databases reveal that information for 'wild' birds is actually greater than that for 'wild' mammals. Of the various taxa of mammals, ecotoxicological data is most noticeably lacking for marsupials and monotremes. In contrast, rodents (comprising 43% of all mammal species) have been studied extensively, despite evidence of their tolerance to some organochlorine compounds, rodenticides, and even radionuclides. Mammalian species at greatest risk of exposure include those that consume a high percentage of their body weight on a daily basis (e.g., shrews, moles and bats). Aquatic mammals tend to bioaccumulate tremendous burdens of lipophilic contaminants, although storage in their fat depots may actually limit toxicity. Carnivores appear to be more sensitive to adverse effects of environmental contaminants than herbivores. Remarkably few of the thousands of compounds manufactured worldwide have been toxicologically evaluated in wild mammals, and concentrations of even fewer have been monitored in tissues. Overarching research needs include: development of new exposure/effects models and better methods for estimation of species sensitivities; generation of comparative data on contaminant bioavailability, sublethal responses and detoxication mechanisms; enhanced understanding of pesticide, industrial contaminant and metal interactions; identification of endocrine disruptive contaminants and their overall ecological significance; and finally, estimating the relative contribution of environmental contamination as a factor affecting wild mammal populations.

Society of Environmental Toxicology and Chemistry,

Masked expression of life-history traits in a highly variable environment

Differing life-history strategies may act as a constraint on reproductive expression that ultimately limits the ability of individual species to respond to changes in the magnitude or frequency of environmental variation, and potentially underlies the variation often inherent in phenotypic and evolved responses to anthropogenic change. Alternatively, if there are environmental cues that predict reproductive potential, differential expression of life-history strategies may represent differences in the adaptive capacity to optimize current reproductive value given variation in environmental conditions. We compared several aspects of walleye Sander vitreus spawning ecology at two reservoirs that differ in environmental variability (i.e., annual water-level fluctuation) to identify the capacity of phenotypic expression and the corresponding association with age. Despite significant differences in female body and liver masses between reservoirs that differ in environmental variability, we found no difference in reproductive investment measured by egg size and fecundity. Walleye in a highly variable environment appear to exhibit reproductive traits more typical of a short-lived life-history strategy, which may be resultant from the interaction of environmental and anthropogenic pressures. This finding emphasizes the need to identify the degree to which life-history expression represents physiological constraints versus ecological optimization, particularly as anthropogenic change continues to alter environmental conditions.

Great Plains Research

Coastal Prairie Restoration Information System: Version 1 (Louisiana)

The Coastal Prairie Restoration Information System (CPR) is a Microsoft Access database that allows users to query and view data about Louisiana coastal prairie species. Less than 0.1% of Louisiana's coastal prairie vegetation remains in a relatively undisturbed condition. Encompassing as much as 1 million hectares of land, coastal prairie is a hybrid of coastal wetlands and tall grass prairie. Over 550 plant species have been identified in Louisiana's coastal prairies to date. Efforts to conserve and restore this endangered ecosystem are limited by the ability of workers to identify and access knowledge about this diverse group of plants. In this database, a variety of data are provided for each of 650 coastal prairie species in Louisiana. The database was developed at the U.S. Geological Survey National Wetlands Research Center by Larry Allain, with software development by Myra Silva. Additional funding was provided by the biology department of the University of Louisiana at Lafayette (ULL), the ULL Center for Environmental and Ecological Technology, and the National Science Foundation.

Data Series

Changes in aquatic prey resources in response to estuary restoration in Willapa Bay, southwestern Washington

Executive Summary The ongoing restoration of more than 200 hectares of estuarine habitat at Willapa National Wildlife Refuge, southwestern Washington, is expected to benefit a variety of species, including salmonids that use estuarine and tidal marshes as rearing and feeding areas as well as migratory waterbirds. During March–June 2014 and 2015, U.S. Geological Survey Western Ecological Research Center (WERC) initiated a study to assess aquatic prey resources, in coordination with a separate but parallel fish study done by the Columbia River Estuary Study Taskforce. WERC collected data on environmental variables and invertebrate community structure, and the taskforce provided salmonid diet data at restored (Lewis Stream and Porter Point) and reference (Greenhead Slough and Ellsworth Creek) sites. We analyzed these data to determine the functional capacity of the estuary for supporting invertebrate prey resources for fish following restoration. The results of our analyses were as follows: Water temperatures were roughly 0.74 degrees Celsius warmer in 2015 than in 2014 at all sites, with potential consequences for salmonid bioenergetics in June and July. Mudflat was colonized by low marsh species such as pickleweed from 2014 to 2015 at restored Lewis Stream. Vegetation community structure remained stable at Greenhead Slough, Ellsworth Creek, and Porter Point in both years, and consisted of halophilic sedges (for example, Carex lyngbyei ) and saltgrass ( Distichlis spicata ). Benthic invertebrate community structure consisted of Polychaeta, Nematoda, Oligochaeta, Amphipoda, and Diptera larvae, all of which contribute to the foraging capacity of juvenile salmon and migratory shorebirds. Benthic invertebrate biomass increased as much as 30-fold at some sites from 2014 to 2015. Terrestrial invertebrate community structure was dominated by Dipteran flies, especially at restored Lewis Stream, which primarily was unvegetated in 2014. Other key taxa included Hemiptera, Arachnida, and Collembola. Aquatic invertebrate prey consisted of planktonic taxa and terrestrial invertebrate drift that fell into the water column from overhanging vegetation. The restored Porter Point had markedly fewer Copepoda, but had the highest levels of neuston biomass primarily due to Dipteran drift (terrestrial flies that fell into the water column). Average proportion similarity index (PSI) values between salmon diet and invertebrate prey availability were relatively low at all sites (<0.1), but were highest at the restored Lewis Stream (0.105±0.102). This likely was influenced by the predominance of Diptera in the surrounding habitat and in the diets of juvenile Chinook and chum salmon ( Oncorhynchus tshawytscha and O. keta , respectively). The invasive New Zealand mud snail ( Potamopyrgus antipodarum ) was detected at both Lewis Stream (8 out of 34 aquatic and benthic samples) and Porter Point (1 out of 32 aquatic and benthic samples) restoration sites, but not at reference sites Greenhead Slough or Ellsworth Creek. This invasive species has been observed throughout the Pacific Northwest coastline is not palatable to most fish (including salmonids) and may even be detrimental to some fish species. Although invertebrate communities differed between restored and reference sites, invertebrate biomass at the restored Lewis Stream and Porter Point was like or exceeded that of reference sites Greenhead Slough and Ellsworth Creek. The restored sites are still in the early phases of restoration and succession, but our study suggests they have the capacity to support foraging wildlife species such as salmonids.

Washington

The ecology of Humboldt Bay, California, an estuarine profile

Synthesizes data on the ecological relationships and functions of the estuary, including information on physical aspects of the bay environment; describes the biotic communities and their relationships; provides management considerations in terms including environmental concerns; and identifies research and management information gaps. Portions of the bay are managed as a national wildlife refuge. Management issues for this ecosystem include loss of habitat and degradation of the environment by additional industrial development and nonpoint source pollution. -from Authors

California

Purpose, processes, partnerships, and products: four Ps to advance participatory socio-environmental modeling

Including stakeholders in environmental model building and analysis is an increasingly popular approach to understanding ecological change. This is because stakeholders often hold valuable knowledge about socio-environmental dynamics and collaborative forms of modeling produce important boundary objects used to collectively reason about environmental problems. Although the number of participatory modeling (PM) case studies and the number of researchers adopting these approaches has grown in recent years, the lack of standardized reporting and limited reproducibility have prevented PM's establishment and advancement as a cohesive field of study. We suggest a four-dimensional framework (4P) that includes reporting on dimensions of (1) the Purpose for selecting a PM approach (the why ); (2) the Process by which the public was involved in model building or evaluation (the how ); (3) the Partnerships formed (the who ); and (4) the Products that resulted from these efforts (the what ). We highlight four case studies that use common PM software-based approaches (fuzzy cognitive mapping, agent-based modeling, system dynamics, and participatory geospatial modeling) to understand human–environment interactions and the consequences of ecological changes, including bushmeat hunting in Tanzania and Cameroon, agricultural production and deforestation in Zambia, and groundwater management in India. We demonstrate how standardizing communication about PM case studies can lead to innovation and new insights about model-based reasoning in support of ecological policy development. We suggest that our 4P framework and reporting approach provides a way for new hypotheses to be identified and tested in the growing field of PM.

Ecological Applications

Impacts to wildlife of wind energy siting and operation in the United States

Electricity from wind energy is a major contributor to the strategy to reduce greenhouse gas emissions from fossil fuel use and thus reduce the negative impacts of climate change. Wind energy, like all power sources, can have adverse impacts on wildlife. After nearly 25 years of focused research, these impacts are much better understood, although uncertainty remains. In this report, we summarize positive impacts of replacing fossil fuels with wind energy, while describing what we have learned and what remains uncertain about negative ecological impacts of the construction and operation of land-based and offshore wind energy on wildlife and wildlife habitat in the U.S. Finally, we propose research on ways to minimize these impacts. TO SUMMARIZE: 1 Environmental and other benefits of wind energy include near-zero greenhouse gas emissions, reductions of other common air pollutants, and little or no water use associated with producing electricity from wind energy. Various scenarios for meeting U.S. carbon emission reduction goals indicate that a four- to five-fold expansion of land-based wind energy from the current 97 gigawatts (GW) by the year 2050 is needed to minimize temperature increases and reduce the risk of climate change to people and wildlife. 2 Collision fatalities of birds and bats are the most visible and measurable impacts of wind energy production. Current estimates suggest most bird species, especially songbirds, are at low risk of population-level impacts. Raptors as a group appear more vulnerable to collisions. Population-level impacts on migratory tree bats are a concern, and better information on population sizes is needed to evaluate potential impacts to these species. Although recorded fatalities of cave-dwelling bat species are typically low at most wind energy facilities, additional mortality from collisions is a concern given major declines in these species due to white-nose syndrome (WNS). Assessments of regional and cumulative fatality impacts for birds and bats have been hampered by the lack of data from areas with a high proportion of the nation’s installed wind energy capacity. Efforts to expand data accessibility from all regions are underway, and this greater access to data along with improvements in statistical estimators should lead to improved impact assessments. 3 Habitat impacts of wind energy development are difficult to assess. An individual wind energy facility may encompass thousands of acres, but only a small percentage of the landscape within the project area is directly transformed. If a project is sited in previously undisturbed habitat, there is concern for indirect impacts, such as displacement of sensitive species. Studies to date indicate displacement of some species, but the long-term population impacts are unknown. 4 Offshore wind energy development in the U.S. is just beginning. Studies at offshore wind facilities in Europe indicate some bird and marine mammal species are displaced from project areas, but substantial uncertainty exists regarding the individual or population-level impacts of this displacement. Bird and bat collisions with offshore turbines are thought to be less common than at terrestrial facilities, but currently the tools to measure fatalities at offshore wind energy facilities are not available. The wind energy industry, state and federal agencies, conservation groups, academia, and scientific organizations have collaborated for nearly 25 years to conduct the research needed to improve our understanding of risk to wildlife and to avoid and minimize that risk. Efforts to reduce the uncertainty about wildlife risk must keep up withthe pace and scale of the need to reduce carbon emissions. This will require focusing our research priorities and increasing the rate at which we incorporate research results into the development and validation of best practices for siting and operating wind energy facilities.

Issues in Ecology

An evaluation of silver-stage American Eel conspecific chemical cueing during outmigration

American Eel Anguilla rostrata abundance has declined in recent decades, in part because sexually maturing, silver-stage adults, outmigrating from freshwater to oceanic spawning grounds, encounter migratory blockades or perish when passing through active hydroelectric turbines. To help improve downstream passage effectiveness and increase survival rates, the role of silver-stage American Eel conspecific chemical cueing during outmigration was investigated using a new type of bioassay. Inside a laboratory flume, downstream-swimming eels were exposed to both live (putative attractant) and dead (putative repellent) conspecific washings to determine whether their trajectory of downstream movement, level of activity, or time spent inside targeted areas of the arena changed after exposure. Silver eels were not attracted to or repulsed by either odor, as none of five scoring metrics indicated a behavioral response. Results did not support the hypothesis that conspecific chemical cueing is a mechanism for downstream migration coordination or danger avoidance; however responses may not have been readily apparent in this type of assay. Fisheries managers may opt to focus future research on more feasible restoration efforts using alternate experimental designs to remedy this ecological issue.

Environmental Biology of Fishes

Using science-policy integration to improve ecosystem science and inform decision-making: Lessons from U.S. LTERs

This Special Session took place on 12 August 2015 at the 100th Meeting of the Ecological Society of America in Baltimore, Maryland, and was conceived of and coordinated by the Science Policy Exchange. The Science Policy Exchange (SPE) is a boundary- spanning organization established to work at the interface of science and policy to confront pressing environmental challenges . SPE was created as a collaborative of six research institutions to increase the impact of science on environmental decisions. This session was organized by Marissa Weiss and co- organized by Pamela Templer, Kathleen Fallon Lambert, Jill Baron, Charles Driscoll, and David Foster. Along the theme of ESA &rsquo; s Centennial meeting, the group of presenters represented collectively more than 100 years of experience in integration of science, policy, and outreach.

Bulletin of the Ecological Society of America

USGS Science Serves Public Health

Human health so often depends on the health of the environment and wildlife around us. The presence of naturally occurring or human environmental contaminants and the emergence of diseases transferred between animals and humans are growing concerns worldwide. The USGS is a source of natural science information vital for understanding the quantity and quality of our earth and living resources. This information improves our understanding not only of how human activities affect environmental and ecological health, but also of how the quality of our environment and wildlife in turn affects human health. USGS is taking a leadership role in providing the natural science information needed by health researchers, policy makers, and the public to safeguard public health

Fact Sheet

A watershed approach to ecosystem monitoring in Denali National Park and preserve, Alaska

The National Park Service and the National Biological Service initiated research in Denali National Park and Preserve, a 2.4 million-hectare park in southcentral Alaska, to develop ecological monitoring protocols for national parks in the Arctic/Subarctic biogeographic area. We are focusing pilot studies on design questions, on scaling issues and regionalization, ecosystem structure and function, indicator selection and evaluation, and monitoring technologies. Rock Creek, a headwater stream near Denali headquarters, is the ecological scale for initial testing of a watershed ecosystem approach. Our conceptual model embraces principles of the hydrological cycle, hypotheses of global climate change, and biological interactions of organisms occupying intermediate, but poorly studied, positions in Alaskan food webs. The field approach includes hydrological and depositional considerations and a suite of integrated measures linking key aquatic and terrestrial biota, environmental variables, or defined ecological processes, in order to establish ecological conditions and detect, track, and understand mechanisms of environmental change. Our sampling activities include corresponding measures of physical, chemical, and biological attributes in four Rock Creek habitats believed characteristic of the greater system diversity of Denali. This paper gives examples of data sets, program integration and scaling, and research needs.

Journal of the American Water Resources Associatio

To predict the niche, model colonization and extinction

Ecologists frequently try to predict the future geographic distributions of species. Most studies assume that the current distribution of a species reflects its environmental requirements (i.e., the species' niche). However, the current distributions of many species are unlikely to be at equilibrium with the current distribution of environmental conditions, both because of ongoing invasions and because the distribution of suitable environmental conditions is always changing. This mismatch between the equilibrium assumptions inherent in many analyses and the disequilibrium conditions in the real world leads to inaccurate predictions of species' geographic distributions and suggests the need for theory and analytical tools that avoid equilibrium assumptions. Here, we develop a general theory of environmental associations during periods of transient dynamics. We show that time-invariant relationships between environmental conditions and rates of local colonization and extinction can produce substantial temporal variation in occupancy&ndash;environment relationships. We then estimate occupancy&ndash;environment relationships during three avian invasions. Changes in occupancy&ndash;environment relationships over time differ among species but are predicted by dynamic occupancy models. Since estimates of the occupancy&ndash;environment relationships themselves are frequently poor predictors of future occupancy patterns, research should increasingly focus on characterizing how rates of local colonization and extinction vary with environmental conditions.

Ecology