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Hydrological connectivity for riverine fish: measurement challenges and research opportunities

In this review, we first summarize how hydrologic connectivity has been studied for riverine fish capable of moving long distances, and then identify research opportunities that have clear conservation significance. Migratory species, such as anadromous salmonids, are good model organisms for understanding ecological connectivity in rivers because the spatial scale over which movements occur among freshwater habitats is large enough to be easily observed with available techniques; they are often economically or culturally valuable with habitats that can be easily fragmented by human activities; and they integrate landscape conditions from multiple surrounding catchment(s) with in‐river conditions. Studies have focussed on three themes: (i) relatively stable connections (connections controlled by processes that act over broad spatio‐temporal scales >1000 km 2 and >100 years); (ii) dynamic connections (connections controlled by processes acting over fine to moderate spatio‐temporal scales ∼1–1000 km 2 and <1–100 years); and (iii) anthropogenic influences on hydrologic connectivity, including actions that disrupt or enhance natural connections experienced by fish. We outline eight challenges to understanding the role of connectivity in riverine fish ecology, organized under three foci: (i) addressing the constraints of river structure; (ii) embracing temporal complexity in hydrologic connectivity; and (iii) managing connectivity for riverine fishes. Challenges include the spatial structure of stream networks, the force and direction of flow, scale‐dependence of connectivity, shifting boundaries, complexity of behaviour and life histories and quantifying anthropogenic influence on connectivity and aligning management goals. As we discuss each challenge, we summarize relevant approaches in the literature and provide additional suggestions for improving research and management of connectivity for riverine fishes. Specifically, we suggest that rapid advances are possible in the following arenas: (i) incorporating network structure and river discharge into analyses; (ii) increasing explicit consideration of temporal complexity and fish behaviour in the scope of analyses; and (iii) parsing degrees of human and natural influences on connectivity and defining acceptable alterations. Multiscale analyses are most likely to identify dominant patterns of connections and disconnections, and the appropriate scale at which to focus conservation activities.

Freshwater Biology

Introduced species as evolutionary traps

Invasive species can alter environments in such a way that normal behavioural decision-making rules of native species are no longer adaptive. The evolutionary trap concept provides a useful framework for predicting and managing the impact of harmful invasive species. We discuss how native species can respond to changes in their selective regime via evolution or learning. We also propose novel management strategies to promote the long-term co-existence of native and introduced species in cases where the eradication of the latter is either economically or biologically unrealistic.

Ecology Letters

Fort Collins Science Center- Policy Analysis and Science Assistance Branch : Integrating social, behavioral, economic and biological sciences

The Fort Collins Science Center's Policy Analysis and Science Assistance (PASA) Branch is a team of approximately 22 scientists, technicians, and graduate student researchers. PASA provides unique capabilities in the U.S. Geological Survey by leading projects that integrate social, behavioral, economic, and biological analyses in the context of human-natural resource interactions. Resource planners, managers, and policymakers in the U.S. Departments of the Interior (DOI) and Agriculture (USDA), State and local agencies, as well as international agencies use information from PASA studies to make informed natural resource management and policy decisions. PASA scientists' primary functions are to conduct both theoretical and applied social science research, provide technical assistance, and offer training to advance performance in policy relevant research areas. Management and research issues associated with human-resource interactions typically occur in a unique context, involve difficult to access populations, require knowledge of both natural/biological science in addition to social science, and require the skill to integrate multiple science disciplines. In response to these difficult contexts, PASA researchers apply traditional and state-of-the-art social science methods drawing from the fields of sociology, demography, economics, political science, communications, social-psychology, and applied industrial organization psychology. Social science methods work in concert with our rangeland/agricultural management, wildlife, ecology, and biology capabilities. The goal of PASA's research is to enhance natural resource management, agency functions, policies, and decision-making. Our research is organized into four broad areas of study.

Fact Sheet

Implementation of the CREED approach for environmental assessments

Environmental exposure data are a key component of chemical and ecological assessments, supporting and guiding environmental management decisions and regulations. Measures taken to protect the environment based on exposure data can have social and economic implications. Flawed information may lead to measures being taken in the wrong place or to important action not being taken. Although the advantages of harmonizing evaluation methods have been demonstrated for hazard information, no comparable approach is established for exposure data evaluation. The goal of Criteria for Reporting and Evaluating Exposure Datasets (CREED) is to improve the transparency and consistency with which exposure data are evaluated regarding usability in environmental assessments. Here, we describe the synthesis of the CREED process, and propose methods and tools to summarize and interpret the outcomes of the data usability evaluation in support of decision-making and communication. The CREED outcome includes a summary that reports any key gaps or shortcomings in the reliability (data quality) and relevance (fitness for purpose) of the data being considered. The approach has been implemented in a workbook template (provided as Supporting Information), for assessors to readily follow the workflow and create a report card for any given dataset. The report card communicates the outcome of the CREED evaluation and summarizes important dataset attributes, providing a concise reference pertaining to the dataset usability for a specified purpose and documenting data limitations that may restrict data use or increase environmental assessment uncertainty. The application of CREED is demonstrated through three case studies, which also were used during beta testing of the methodology. As experience with the CREED approach application develops, further improvements may be identified and incorporated into the framework. Such development is to be encouraged in the interest of better science and decision-making, and to make environmental monitoring and assessment more cost-effective.

Integrated Environmental Assessment and Management

The role of habitat filtering in the leaf economics spectrum and plant susceptibility to pathogen infection

1.The Leaf Economics Spectrum (LES) describes global covariation in the traits of plant leaves. The LES is thought to arise from biophysical constraints and habitat filtering (ecological selection against unfit trait combinations along environmental gradients). However, the role of habitat filtering in generating the LES has not been tested experimentally. 2.If the process of habitat filtering plays a role in generating the LES, the LES could weaken in communities that have yet to be filtered by the current environment, for example after abiotic environmental change. LES traits are commonly used to predict community and ecosystem processes, and if the LES weakens in unfiltered communities, LES-based models may no longer apply. 3.In the greenhouse, we experimentally simulated three stages of habitat filtering in response to abiotic change: from unfiltered, to semi-filtered, to completely filtered communities. In each stage, we quantified the strength of the LES and assessed the accuracy of trait-based models of an important ecological process, pathogen infection. 4.The strength of the LES increased with the completeness of habitat filtering, as did the accuracy of trait-based models of plant susceptibility to pathogen infection. 5. Synthesis . Our results suggest that habitat filtering plays a fundamental role in strengthening the trait correlations of the LES, and that trait-based models may be less accurate when communities have not been filtered by the current environment, for example, following rapid environmental change.

Journal of Ecology

Infection by the marine cestode Hepatoxylon trichiuri in returning Chinook salmon (Oncorhynchus tshawytscha) spawners in Patagonia: Implications for a novel fishery

Parasitological information may be useful for managing economically important fisheries by providing insights into population dynamics and health effects. The Chinook salmon ( Oncorhynchus tshawytscha ), an invasive species with high socio-ecological importance in South America, supports recreational and incipient artisanal fisheries. In Southern Chile, the recent approval of coastal fisheries for returning Chinook salmon in some regions highlights the importance of understanding salmon foraging, habitat use, and commercial value. However, such information is lacking. Parasites have been extensively used as cost-effective biological tags to identify fish stocks. We conducted the first parasitological examination of the marine tapeworm Hepatoxylon trichiuri larvae in returning adult Chinook salmon from the Cisnes River to assess its potential as a biological tag for returning Chinook salmon in Patagonia. Of 61 Chinook salmon from the Cisnes River, 52.46% were infected by H. trichiuri , with 1.65 ± 1.26 parasites per infected salmon (mean ± SD). Infection prevalence and intensity did not differ significantly between males and females. Infection intensity was higher in larger salmon but decreased with increasing salmon condition. Infection intensity was not significantly related to length, mass, or body condition. Infection prevalence and intensity of H. trichiuri varied spatially among Patagonian river-basin populations of Chinook salmon, thereby supporting the use of H. trichiuri as a biological tag. Our findings provide important parasitological metrics that may enhance future management practices by informing geographic variability in foraging locations of adult salmon and their commercial value for human consumption.

Cisnes River, Patagonia

C4 photosynthesis, trait spectra, and the fast-efficient phenotype

It has been 60 years since the discovery of C 4 photosynthesis, an event that rewrote our understanding of plant adaptation, ecosystem responses to global change, and global food security. Despite six decades of research, one aspect of C 4 photosynthesis that remains poorly understood is how the pathway fits into the broader context of adaptive trait spectra, which form our modern view of functional trait ecology. The C 4 CO 2 -concentrating mechanism supports a general C 4 plant phenotype capable of fast growth and high resource-use efficiencies. The fast-efficient C 4 phenotype has the potential to operate at high productivity rates, while allowing for less biomass allocation to root production and nutrient acquisition, thereby providing opportunities for the evolution of novel trait covariances and the exploitation of new ecological niches. We propose the placement of the C 4 fast-efficient phenotype near the acquisitive pole of the world-wide leaf economic spectrum, but with a pathway-specific span of trait space, wherein selection shapes both acquisitive and conservative adaptive strategies. A trait-based perspective of C 4 photosynthesis will open new paths to crop improvement, global biogeochemical modeling, the management of invasive species, and the restoration of disturbed ecosystems, particularly in grasslands.

New Phytologist

Visitor use and activities detected using trail cameras at forest restoration sites

We used trail cameras to monitor human visits and activities at two sites in northeast Indiana being restored to bottomland hardwood forests. These sites, managed as nature preserves, are close to cities, where trails and parking lots have been added for ease of access. In this study, trail cameras were successfully used to capture visitation rates and activity types. The two sites had median visitor use rates of 1 and 13 visitors per day. Across both sites, “parking lot use only” (62%), hikers (30.2%), and bicyclists (5%) accounted for more than 97% of site visits. Overall, most weekday visitor-time occurred during daylight hours, peaking at lunch and evening. Mean total number of daily visitors was higher during weekends; however, total daily visitor-time did not vary between days of the week. Michaelis-Menten rarefaction models of sampling efficiency across the study’s four camera stations suggest sampling duration of 27 to 55 days to accurately estimate mean daily visitor counts and 3 to 40 days to detect half the maximal numbers of observed activities. Study estimates of visitation provide land managers with information for accommodating visitor use activities on the restored sites and offer inputs for cultural ecosystem services assessments and associated economic analyses.

Ecological Restoration

A typology of time-scale mismatches and behavioral interventions to diagnose and solve conservation problems

Ecological systems often operate on time scales significantly longer or shorter than the time scales typical of human decision making, which causes substantial difficulty for conservation and management in socioecological systems. For example, invasive species may move faster than humans can diagnose problems and initiate solutions, and climate systems may exhibit long-term inertia and short-term fluctuations that obscure learning about the efficacy of management efforts in many ecological systems. We adopted a management-decision framework that distinguishes decision makers within public institutions from individual actors within the social system, calls attention to the ways socioecological systems respond to decision makers’ actions, and notes institutional learning that accrues from observing these responses. We used this framework, along with insights from bedeviling conservation problems, to create a typology that identifies problematic time-scale mismatches occurring between individual decision makers in public institutions and between individual actors in the social or ecological system. We also considered solutions that involve modifying human perception and behavior at the individual level as a means of resolving these problematic mismatches. The potential solutions are derived from the behavioral economics and psychology literature on temporal challenges in decision making, such as the human tendency to discount future outcomes at irrationally high rates. These solutions range from framing environmental decisions to enhance the salience of long-term consequences, to using structured decision processes that make time scales of actions and consequences more explicit, to structural solutions aimed at altering the consequences of short-sighted behavior to make it less appealing. Additional application of these tools and long-term evaluation measures that assess not just behavioral changes but also associated changes in ecological systems are needed.

Conservation Biology

Fishing for conservation of freshwater tropical fish in the Anthropocene

1. Biodiversity and fisheries are two important assets of freshwater ecosystems that are currently at risk from external threats. Establishing an equitable resolution to these threats is a major challenge of the Anthropocene. 2. This is particularly pertinent in developing nations where hotspots for biodiversity converge with rapid, and often environmentally degrading, economic development, and high dependency on aquatic biota and fisheries by local communities. 3. Here, we present global case studies that demonstrate how building upon a shared need for healthy environments and ecological integrity can meet both biodiversity conservation and fishery objectives. 4. These case studies provide evidence that addressing biodiversity conservation needs is possible through partnerships (especially with fishers), shared knowledge, and innovation in fisheries management. 5. In the pursuit of sustainable use of fresh water in development, both conservation and fisheries agendas are better served if efforts focused on synergising fishing activities with local ecosystem functioning and yield long-term livelihood and food security perspectives. 6. A partnership between conservationists (e.g., practitioners, environmental NGOs) and fisheries communities provides a unified voice advocating for biodiversity and social interests in freshwater governance decisions, which has more socio-economic and political capital than when voices are independent or in competition.

Aquatic Conservation: Marine and Freshwater Ecosys

Research plan for lands administered by the U.S. Department of the Interior in the Interior Columbia Basin and Snake River Plateau

This document presents a long-term research strategy designed to address current and future research needs for management of Department of the Interior-administered ecosystems in the Intermountain West. Although the research plan was developed in the context of the Interior Columbia Basin Ecosystem Management Project, the plan addresses many high-priority issues facing land managers throughout the Intermountain West. These issues pose management challenges that may be addressed with applied research both currently and in upcoming decades. Possessing a particular focus on semiarid ecosystems, the plan is a collection of research questions under five categories of research emphases: 1) restoration; 2) rangeland health; 3) aquatic-terrestrial connections; 4) development of monitoring and evaluation protocols; and 5) species and habitats at risk. The goal of the research strategy is to provide ideas for integrating emerging scientific understanding into future management in order to restore and maintain long-term ecosystem health and ecological integrity; provide consistent management direction over broad spatial and temporal scales; emphasize adaptive management over the long term; restore and maintain habitats for plant and animal species; and support economic and social needs of people, without compromising the above goals. Research questions are prioritized into three categories based on the immediacy of their need, feasibility of addressing the question rigorously under varying funding budgets, and magnitude of risk posed by not addressing the issue. The research strategy is intended to support and integrate with existing management efforts and strategies. As such, it melds observational studies with experimental manipulation, treating management actions as experiments whenever feasible. The research strategy focuses on disturbance processes and events that have been the primary drivers of change, to provide a predictive model for future changes. These drivers include fire, nonnative plants, herbivory, roads and associated human influences, and climate change. Whereas management in the western United States has striven to move from an inefficient species-based approach to a habitat-based approach, the plan focuses on ecosystem function and ecological processes as critical measures of habitat response. Because of the large amount and contiguity of public lands in the western United States, the region presents both a compelling opportunity to implement landscape-level science and a challenge to underst

Information and Technology Report

Temporal habitat use of mule deer in the Pueblo of Santa Ana, New Mexico

Mule deer ( Odocoileus hemionus ) are important economically, culturally, and recreationally to the Pueblo of Santa Ana in central New Mexico, USA. Studies of habitat selection improve our understanding of mule deer ecology in central New Mexico and provide the Tribe with valuable information for management of mule deer. We used global positioning system telemetry-collar data collected on mule deer around the Pueblo of Santa Ana to create resource selection functions from proximity-based habitat predictors using a generalized linear mixed model. We created separate resource selection functions for females and males during summer and winter at different times of the day. Season generally had a greater effect on mule deer habitat use than the time of day. Female and male mule deer selected for similar habitats but were sexually segregated in their summer distributions. These findings are consistent with results from other locations where mule deer partitioned habitat similarly between seasons and sexes. Supported models reaffirm accepted patterns of habitat selection for mule deer to the Pueblo of Santa Ana where local results were lacking. Our results can help managers identify locations in and around the Pueblo of Santa Ana where future development such as highway expansion are likely to conflict with mule deer activity and locations where habitat enhancement projects such as adding water sources can have the greatest effect for the deer population.

New Mexico

Temporal variability in irrigated land and climate influences on salinity loading across the Upper Colorado River Basin, 1986-2017

Freshwater salinization is a growing global concern impacting human and ecosystem needs with impacts to water availability for human and ecological uses. In the Upper Colorado River Basin (UCRB), dissolved solids in streams compound ongoing water supply challenges to further limit water availability and cause economic damages. Much effort has been dedicated to understanding dissolved solid sources, transport, and management in the region, yet temporal variability in loading from key sources such as irrigated lands and the influence of climate on dissolved solids loading remains unknown. Quantifying the contributions and temporal variability of dissolved solids loads from irrigated lands may benefit salinity management efforts. This study applies a time-varying (dynamic) modeling approach to predict annual dissolved solids loads across the UCRB from 1986 through 2017. Between 66% and 82% of the total accumulated dissolved solids load in the basin is from groundwater (storage and baseflow). Our findings link climate, irrigation, and groundwater, and confirm large storage contributions that have declined slightly with time. Dissolved solids loads increase during wet periods and decrease during dry periods, although the relative contributions of different sources vary little with time. Irrigation enhances loading efficiency relative to unirrigated areas through runoff and groundwater, and can locally be a major source of dissolved solids where irrigation occurs. Results indicate that loads from irrigated areas increase when irrigated area and/or water available for runoff increase. Increased regional aridification over the study period may have contributed to decreasing stream salinity through both quicker surface runoff and lagged groundwater storage processes. Study results may be relevant to salinity management in arid environments where water availability is limited and where irrigation influences salinity loading to streams.

Upper Colorado River Basin

A geologic and anthropogenic journey from the Precambrian to the new energy economy through the San Juan volcanic field

The San Juan volcanic field comprises 25,000 km 2 of intermediate composition mid-Tertiary volcanic rocks and dacitic to rhyolitic calderas including the San Juan&ndash;Uncompahgre and La Garita caldera-forming super-volcanoes. The region is famous for the geological, ecological, hydrological, archeological, and climatological diversity. These characteristics supported ancestral Puebloan populations. The area is also important for its mineral wealth that once fueled local economic vitality. Today, mitigating and/or investigating the impacts of mining and establishing the region as a climate base station are the focuses of ongoing research. Studies include advanced water treatment, the acid neutralizing capacity (ANC) of propylitic bedrock for use in mine-lands cleanup, and the use of soil amendments including biochar from beetle-kill pines. Biochar aids soil productivity and revegetation by incorporation into soils to improve moisture retention, reduce erosion, and support the natural terrestrial carbon sequestration (NTS) potential of volcanic soils to help offset atmospheric CO 2 emissions. This field trip will examine the volcano-tectonic and cultural history of the San Juan volcanic field as well as its geologic structures, economic mineral deposits and impacts, recent mitigation measures, and associated climate research. Field trip stops will include a visit to (1) the Summitville Superfund site to explore quartz alunite-Au mineralization, and associated alteration and new water-quality mitigation strategies; (2) the historic Creede epithermal-polymetallic&ndash;vein district with remarkably preserved resurgent calderas, keystone-graben, and moat sediments; (3) the historic mining town of Silverton located in the nested San Juan&ndash;Silverton caldera complex that exhibits base-metal Au-Ag mineralization; and (4) the site of ANC and NTS studies. En route back to Denver, we will traverse Grand Mesa, a high NTS area with Neogene basalt-derived soils and will enjoy a soak in the geothermal waters of the Aspen anomaly at Glenwood Springs.

Colorado

A new map of global ecological land units — An ecophysiographic stratification approach

In response to the need and an intergovernmental commission for a high resolution and data-derived global ecosystem map, land surface elements of global ecological pattern were characterized in an ecophysiographic stratification of the planet. The stratification produced 3,923 terrestrial ecological land units (ELUs) at a base resolution of 250 meters. The ELUs were derived from data on land surface features in a three step approach. The first step involved acquiring or developing four global raster datalayers representing the primary components of ecosystem structure: bioclimate, landform, lithology, and land cover. These datasets generally represent the most accurate, current, globally comprehensive, and finest spatial and thematic resolution data available for each of the four inputs. The second step involved a spatial combination of the four inputs into a single, new integrated raster dataset where every cell represents a combination of values from the bioclimate, landforms, lithology, and land cover datalayers. This foundational global raster datalayer, called ecological facets (EFs), contains 47,650 unique combinations of the four inputs. The third step involved an aggregation of the EFs into the 3,923 ELUs. This subdivision of the Earth’s surface into relatively fine, ecological land areas is designed to be useful for various types of ecosystem research and management applications, including assessments of climate change impacts to ecosystems, economic and non-economic valuation of ecosystem services, and conservation planning.

Book

Assessing rates of forest change and fragmentation in Alabama, USA, using the vegetation change tracker model

Forest change is of great concern for land use decision makers and conservation communities. Quantitative and spatial forest change information is critical for addressing many pressing issues, including global climate change, carbon budgets, and sustainability. In this study, our analysis focuses on the differences in geospatial patterns and their changes between federal forests and nonfederal forests in Alabama over the time period 1987–2005, by interpreting 163 Landsat Thematic Mapper (TM) scenes using a vegetation change tracker (VCT) model. Our analysis revealed that for the most part of 1990 s and between 2000 and 2005, Alabama lost about 2% of its forest on an annual basis due to disturbances, but much of the losses were balanced by forest regeneration from previous disturbances. The disturbance maps revealed that federal forests were reasonably well protected, with the fragmentation remaining relatively stable over time. In contrast, nonfederal forests, which are predominant in area share (about 95%), were heavily disturbed, clearly demonstrating decreasing levels of fragmentation during the time period 1987–1993 giving way to a subsequent accelerating fragmentation during the time period 1994–2005. Additionally, the identification of the statistical relationships between forest fragmentation status and forest loss rate and forest net change rate in relation to land ownership implied the distinct differences in forest cutting rate and cutting patterns between federal forests and nonfederal forests. The forest spatial change information derived from the model has provided valuable insights regarding regional forest management practices and disturbance regimes, which are closely associated with regional economics and environmental concerns.

Forest Ecology and Management

Accounting for sampling patterns reverses the relative importance of trade and climate for the global sharing of exotic plants

Aim The distributions of exotic species reflect patterns of human-mediated dispersal, species climatic tolerances and a suite of other biotic and abiotic factors. The relative importance of each of these factors will shape how the spread of exotic species is affected by ongoing economic globalization and climate change. However, patterns of trade may be correlated with variation in scientific sampling effort globally, potentially confounding studies that do not account for sampling patterns. Location Global. Time period Museum records, generally from the 1800s up to 2015. Major taxa studied Plant species exotic to the United States. Methods We used data from the Global Biodiversity Information Facility (GBIF) to summarize the number of plant species with exotic occurrences in the United States that also occur in each other country world-wide. We assessed the relative importance of trade and climatic similarity for explaining variation in the number of shared species while evaluating several methods to account for variation in sampling effort among countries. Results Accounting for variation in sampling effort reversed the relative importance of trade and climate for explaining numbers of shared species. Trade was strongly correlated with numbers of shared U.S. exotic plants between the United States and other countries before, but not after, accounting for sampling variation among countries. Conversely, accounting for sampling effort strengthened the relationship between climatic similarity and species sharing. Using the number of records as a measure of sampling effort provided a straightforward approach for the analysis of occurrence data, whereas species richness estimators and rarefaction were less effective at removing sampling bias. Main conclusions Our work provides support for broad-scale climatic limitation on the distributions of exotic species, illustrates the need to account for variation in sampling effort in large biodiversity databases, and highlights the difficulty in inferring causal links between the economic drivers of invasion and global patterns of exotic species occurrence.

Global Ecology and Biogeography

Northern forest winters have lost cold, snowy conditions that are important for ecosystems and human communities

Winter is an understudied but key period for the socio-ecological systems of northeastern North American forests. A growing awareness of the importance of the winter season to forest ecosystems and surrounding communities has inspired several decades of research, both across the northern forest and at other mid- and high-latitude ecosystems around the globe. Despite these efforts, we lack a synthetic understanding of how winter climate change may impact hydrological and biogeochemical processes and the social and economic activities they support. Here we take advantage of 100 years of meteorological observations across the northern forest region of the northeastern U.S. and eastern Canada to develop a suite of indicators that enable a cross-cutting understanding of 1) how winter temperatures and snow cover have been changing and 2) how these shifts may impact both ecosystems and surrounding human communities. We show that cold and snow-covered conditions have generally decreased over the past 100 years. These trends suggest positive outcomes for tree health as related to reduced fine root mortality and nutrient loss associated with winter frost but negative outcomes as related to the northward advancement and proliferation of forest insect pests. In addition to effects on vegetation, reductions in cold temperatures and snow cover are likely to have negative impacts on the ecology of the northern forest through impacts on water, soils, and wildlife. The overall loss of coldness and snow cover may also have negative consequences for logging and forest products, vector-borne diseases and human health, recreation and tourism, and cultural practices, which together represent important social and economic dimensions for the northern forest region. These findings advance our understanding of how our changing winters may transform the socio- ecological system of a region that has been defined by the contrasting rhythm of the seasons. Our research also identifies a trajectory of change that informs our expectations for the future as the climate continues to warm.

Ecological Applications