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Coast to coast: High genomic connectivity in North American scoters

Dispersal shapes demographic processes and therefore is fundamental to understanding biological, ecological, and evolutionary processes acting within populations. However, assessing population connectivity in scoters ( Melanitta sp.) is challenging as these species have large spatial distributions that span remote landscapes, have varying nesting distributions (disjunct vs. continuous), exhibit unknown levels of dispersal, and vary in the timing of the formation of pair bonds (winter vs. fall/spring migration) that may influence the distribution of genetic diversity. Here, we used double‐digest restriction‐associated DNA sequence (ddRAD) and microsatellite genotype data to assess population structure within the three North American species of scoter (black scoter, M. americana ; white‐winged scoter, M. deglandi ; surf scoter, M. perspicillata ), and between their European congeners (common scoter, M. nigra ; velvet scoter, M. fusca ). We uncovered no or weak genomic structure (ddRAD Φ ST < 0.019; microsatellite F ST < 0.004) within North America but high levels of structure among European congeners (ddRAD Φ ST > 0.155, microsatellite F ST > 0.086). The pattern of limited genomic structure within North America is shared with other sea duck species and is often attributed to male‐biased dispersal. Further, migratory tendencies (east vs. west) of female surf and white‐winged scoters in central Canada are known to vary across years, providing additional opportunities for intracontinental dispersal and a mechanism for the maintenance of genomic connectivity across North America. In contrast, the black scoter had relatively elevated levels of divergence between Alaska and Atlantic sites and a second genetic cluster found in Alaska at ddRAD loci was concordant with its disjunct breeding distribution suggestive of a dispersal barrier (behavioral or physical). Although scoter populations appear to be connected through a dispersal network, a small percentage (<4%) of ddRAD loci had elevated divergence which may be useful in linking areas (nesting, molting, staging, and wintering) throughout the annual cycle.

North America

Soil water availability shapes species richness in mid-latitude shrub steppe plant communities

Questions Ecological communities are controlled by multiple, interacting abiotic and biotic factors that influence the distribution, abundance, and diversity of species. These processes jointly determine resource availability, resource competition, and ultimately species richness. For many terrestrial ecosystems in dryland climates, soil water availability is the most frequent limiting resource for plant species. We used field sampling coupled with process‐based soil water balance modeling to explore the relative importance of multiple macroclimatic, ecohydrological, and biotic variables on plant species and functional type richness at the landscape scale in dryland plant communities. Location Dryland plant communities dominated by big sagebrush ( Artemisia tridentata ) that span climatic and elevational gradients in Wyoming, USA. Methods We quantified species richness at 1,000 m 2 and used multiple regression to determine whether mean climatic conditions, multiple metrics of soil moisture from a soil water balance model (SOILWAT2), soil physical and chemical properties, and shrub stand structure (biotic) variables were related to species and functional type richness. Results Species richness varied between 16 and 54 across sites. We found that species and functional type richness were related to both macroclimate and ecohydrology, but ecohydrology explained slightly more variation than climate. Biotic variables were always secondary to macroclimate and ecohydrology in our models. Variance partitioning revealed that large portions of variability in species (~54%), forb (~47%), and grass (~40%) richness were explained by ecohydrological variables. Conclusions Our results highlight the importance of the spatial and temporal distribution of soil water for dryland plant species richness and suggest that documenting the ways in which climate, vegetation, and soil properties interact to determine soil water availability is critical for understanding biodiversity patterns in dryland plant communities. This work has relevance for other mid‐latitude, shrub‐dominated dryland plant communities where soil water availability strongly influences ecosystem structure and function.

Wyoming

Climatic controls on the global distribution, abundance, and species richness of mangrove forests

Mangrove forests are highly productive tidal saline wetland ecosystems found along sheltered tropical and subtropical coasts. Ecologists have long assumed that climatic drivers (i.e., temperature and rainfall regimes) govern the global distribution, structure, and function of mangrove forests. However, data constraints have hindered the quantification of direct climate-mangrove linkages in many parts of the world. Recently, the quality and availability of global-scale climate and mangrove data have been improving. Here, we used these data to better understand the influence of air temperature and rainfall regimes upon the distribution, abundance, and species richness of mangrove forests. Although our analyses identify global-scale relationships and thresholds, we show that the influence of climatic drivers is best characterized via regional range limit-specific analyses. We quantified climatic controls across targeted gradients in temperature and/or rainfall within 14 mangrove distributional range limits. Climatic thresholds for mangrove presence, abundance, and species richness differed among the 14 studied range limits. We identified minimum temperature-based thresholds for range limits in eastern North America, eastern Australia, New Zealand, eastern Asia, eastern South America, and southeast Africa. We identified rainfall-based thresholds for range limits in western North America, western Gulf of Mexico, western South America, western Australia, Middle East, northwest Africa, east central Africa, and west central Africa. Our results show that in certain range limits (e.g., eastern North America, western Gulf of Mexico, eastern Asia), winter air temperature extremes play an especially important role. We conclude that rainfall and temperature regimes are both important in western North America, western Gulf of Mexico, and western Australia. With climate change, alterations in temperature and rainfall regimes will affect the global distribution, abundance, and diversity of mangrove forests. In general, warmer winter temperatures are expected to allow mangroves to expand poleward at the expense of salt marshes. However, dispersal and habitat availability constraints may hinder expansion near certain range limits. Along arid and semi-arid coasts, decreases or increases in rainfall are expected to lead to mangrove contraction or expansion, respectively. Collectively, our analyses quantify climate-mangrove linkages and improve our understanding of the expected global- and regional-scale effects of climate change upon mangrove forests.

Ecological Monographs

Defining population structure and genetic signatures of decline in the giant garter snake ( Thamnophis gigas ): implications for conserving threatened species within highly altered landscapes

Anthropogenic habitat fragmentation can disrupt the ability of species to disperse across landscapes, which can alter the levels and distribution of genetic diversity within populations and negatively impact long-term viability. The giant gartersnake ( Thamnophis gigas ) is a state and federally threatened species that historically occurred in the wetland habitats of California&rsquo;s Great Central Valley. Despite the loss of 93 % of historic wetlands throughout the Central Valley, giant gartersnakes continue to persist in relatively small, isolated patches of highly modified agricultural wetlands. Gathering information regarding genetic diversity and effective population size represents an essential component for conservation management programs aimed at this species. Previous mitochondrial sequence studies have revealed historical patterns of differentiation, yet little is known about contemporary population structure and diversity. On the basis of 15 microsatellite loci, we estimate population structure and compare indices of genetic diversity among populations spanning seven drainage basins within the Central Valley. We sought to understand how habitat loss may have affected genetic differentiation, genetic diversity and effective population size, and what these patterns suggest in terms of management and restoration actions. We recovered five genetic clusters that were consistent with regional drainage basins, although three northern basins within the Sacramento Valley formed a single genetic cluster. Our results show that northern drainage basin populations have higher connectivity than among central and southern basins populations, and that greater differentiation exists among the more geographically isolated populations in the central and southern portion of the species&rsquo; range. Genetic diversity measures among basins were significantly different, and were generally lower in southern basin populations. Levels of inbreeding and evidence of population bottlenecks were detected in about half the populations we sampled, and effective population size estimates were well below recommended minimum thresholds to avoid inbreeding. Efforts focused on maintaining and enhancing existing wetlands to facilitate dispersal between basins and increase local effective population sizes may be critical for these otherwise isolated populations.

Conservation Genetics

Revisiting the historic distribution and habitats of the Whooping Crane

The endangered Whooping Crane ( Grus americana ) historically had a wide distribution that covered diverse ecoregions across North America while retaining consistent habitat preferences within each ecoregion. We reevaluate the historic information compiled by Robert Porter Allen in 1952 and added 74 other records. Based on the ecological features of historic locations relative to crane life history, we revisit Allen’s description of the whooping crane’s niche and identify four features common to breeding and wintering areas: (1) gentle to rolling topography with an interspersion of wetland and prairie habitats, and relatively sparse cover of trees and shrubs; (2) high densities of shallow, open wetlands or wetland complexes; (3) hydrological regimes that provide reliable conditions for nesting, brood rearing, and flightless adults; and (4) high plant and animal productivity due to fertile soils, hydrological pulsing, periodic inflow of nutrients, or other periodic perturbations. Accurate determination of the ecological features that compose Whooping Crane habitats should stimulate renewed discussions about habitat requirements and can support development of improved reintroduction strategies for the long-term success of recovery efforts.

Book chapter

Genetic population substructure in bison at Yellowstone National Park

The Yellowstone National Park bison herd is 1 of only 2 populations known to have continually persisted on their current landscape since pre-Columbian times. Over the last century, the census size of this herd has fluctuated from around 100 individuals to over 3000 animals. Previous studies involving radiotelemetry, tooth wear, and parturition timing provide evidence of at least 2 distinct groups of bison within Yellowstone National Park. To better understand the biology of Yellowstone bison, we investigated the potential for limited gene flow across this population using multilocus Bayesian clustering analysis. Two genetically distinct and clearly defined subpopulations were identified based on both genotypic diversity and allelic distributions. Genetic cluster assignments were highly correlated with sampling locations for a subgroup of live capture individuals. Furthermore, a comparison of the cluster assignments to the 2 principle winter cull sites revealed critical differences in migration patterns across years. The 2 Yellowstone subpopulations display levels of differentiation that are only slightly less than that between populations which have been geographically and reproductively isolated for over 40 years. The identification of cryptic population subdivision and genetic differentiation of this magnitude highlights the importance of this biological phenomenon in the management of wildlife species.

Journal of Heredity

Evaluating multi-level models to test occupancy state responses of Plethodontid salamanders

Plethodontid salamanders are diverse and widely distributed taxa and play critical roles in ecosystem processes. Due to salamander use of structurally complex habitats, and because only a portion of a population is available for sampling, evaluation of sampling designs and estimators is critical to provide strong inference about Plethodontid ecology and responses to conservation and management activities. We conducted a simulation study to evaluate the effectiveness of multi-scale and hierarchical single-scale occupancy models in the context of a Before-After Control-Impact (BACI) experimental design with multiple levels of sampling. Also, we fit the hierarchical single-scale model to empirical data collected for Oregon slender and Ensatina salamanders across two years on 66 forest stands in the Cascade Range, Oregon, USA. All models were fit within a Bayesian framework. Estimator precision in both models improved with increasing numbers of primary and secondary sampling units, underscoring the potential gains accrued when adding secondary sampling units. Both models showed evidence of estimator bias at low detection probabilities and low sample sizes; this problem was particularly acute for the multi-scale model. Our results suggested that sufficient sample sizes at both the primary and secondary sampling levels could ameliorate this issue. Empirical data indicated Oregon slender salamander occupancy was associated strongly with the amount of coarse woody debris (posterior mean = 0.74; SD = 0.24); Ensatina occupancy was not associated with amount of coarse woody debris (posterior mean = -0.01; SD = 0.29) . Our simulation results indicate that either model is suitable for use in an experimental study of Plethodontid salamanders provided that sample sizes are sufficiently large. However, hierarchical single-scale and multi-scale models describe different processes and estimate different parameters. As a result, we recommend careful consideration of study questions and objectives prior to sampling data and fitting models.

PLoS ONE

Genetics of Central Valley, O. mykiss, populations: Drainage and watershed scale analyses

Genetic variation at 11 microsatellite loci described population genetic structure for Oncorhynchus mykiss in the Central Valley, California. Spatial and temporal variation was examined as well as relationships between hatchery and putative natural spawning anadromous stocks. Genetic diversity was analyzed at two distinct spatial scales: fine-scale within drainage for five populations on Clear Creek; between and among drainage diversity for 23 populations. Significant regional spatial structure was apparent, both within Clear Creek and among rainbow trout populations throughout the Central Valley. Significant differences in allelic frequencies were found among most river or drainage systems. Less than 1% of the molecular variance could be attributed to differences found between drainages. Hatchery populations were shown to carry similar genetic diversity to geographically proximate wild populations. Central Valley M = 0.626 (below the M < 0.68 threshold) supported recent population reductions within the Central Valley. However, average estimated effective population size was relatively high (Ne = 5066). Significant allelic differences were found in rainbow trout collected above and below impassable dams on the American, Yuba, Stanislaus and Tuolumne rivers. Rainbow trout sampled in Spring Creek were extremely bottlenecked with allelic variation at only two loci and an estimated effective population size of 62, suggesting some local freshwater O. mykiss stocks may be declining rapidly. These data support significant genetic population structure for steelhead and rainbow trout populations within the Central Valley across multiple scales. Careful consideration of this genetic diversity and its distribution across the landscape should be part of future conservation and restoration efforts.

California

Assessing the role of seabirds in the ecology of influenza A viruses

Wild waterbirds, specifically waterfowl, gulls, and shorebirds, are recognized as the primordial reservoir of influenza A viruses (IAVs). However, the role of seabirds, an abundant, diverse, and globally distributed group of birds, in the perpetuation and transmission of IAVs is less clear. Here we summarize published and publicly available data for influenza viruses in seabirds, which for the purposes of this study are defined as birds that exhibit a largely or exclusively pelagic lifestyle and exclude waterfowl, gulls, and shorebirds, and we review this collective dataset to assess the role of seabirds in the influenza A ecology. Since 1961, more than 40,000 samples have been collected worldwide from the seabirds considered here and screened, using a variety of techniques, for evidence of active or past IAV infection. From these data, the overall prevalence of active infection has been estimated to be very low; however, serological data provide evidence that some seabird species are more frequently exposed to IAVs. Sequence data for viruses from seabirds are limited, except for murres (common murre, Uria aalge , and thick-billed murre, Uria lomvia ; family Alcidae) for which there are full or partial genome sequences available for more than 80 viruses. Characterization of these viruses suggests that murres are infected with Group 1 hemagglutinin subtype viruses more frequently as compared to Group 2 and also indicates that these northern, circumpolar birds are frequently infected by intercontinental reassortant viruses. Greater temporal and spatial sampling and characterization of additional viruses are required to better understand the role of seabirds in global IAV dynamics.

Avian Diseases

Surface-water-quality assessment of the upper Illinois River basin in Illinois, Indiana, and Wisconsin; project description

In 1986, the U.S. Geological Survey began a National Water-Quality Assessment program to (1) provide nationally consistent descriptions of the current status of water quality for a large, diverse, and geographically distributed part of the Nation's surface- and ground-water resources; (2) define, where possible, trends in water quality; and (3) identify and describe the relations of both status and trends in water quality to natural factors and the history of land use and land- and waste-management activities. The program is presently in a pilot phase that will test and modify, as necessary, concepts and approaches in preparation for possible full implementation of the program in the future. The upper Illinois River basin is one of four basins selected to test the concepts and approaches of the surface-water-quality element of the national program. The basin drains 10,949 square miles of Illinois, Indiana, and Wisconsin. Three principal tributaries are the Kankakee and Des Plaines Rivers that join to form the Illinois River and the Fox River. Land use is predominantly agricultural; about 75 percent of the basin is cultivated primarily for production of corn and soybeans. About 13 percent of the basin is urban area, most of which is located in the Chicago metropolitan area. The population of the basin is about 7 million. About 6 million people live in the Des Plaines River basin. Many water-quality issues in the upper Illinois River basin are related to sediment, nutrients, potentially toxic inorganic and organic constituents, and to water-management practices. Occurrence of sediment and the chemical constituents in the rivers and lakes within the basin has the potential to adversely affect the water's suitability for aquatic life, recreation, or, through the consumption of fish, human health. The upper Illinois River basin project consists of five major activities. The first activity--analysis of existing information and preparation of a report that describes recent water-quality conditions and trends--is currently underway. The second activity--fixed-station water-quality sampling at eight stations--began in April 1987 and will last at least 3 years. Water-quality data collected at these stations will be used to determine the frequency of occurrence of constituent concentrations, their annual and seasonal loads, and time trends in concentrations for a selected number of constituents. The third activity will be synoptic water-quality studies. Each study will involve sampling many sites at specific flow conditions and for selected water-quality constituents. Information gained from these studies will supplement informa tion gained from fixed-station sampling. A synoptic study of streambed sediments is tentatively planned for the summer of 1987 to describe the occurrence and distribution of trace elements in the basin. The fourth activity will consist of one or more topical subbasin or river-reach studies. The purpose of such studies is to better define certain water-quality conditions in specific areas and gain an understanding of the processes affecting the observed conditions. The fifth activity is the preparation of reports that will describe results from each of the first four activities. Quality assurance and coordination are being provided at both the national and pilot-project levels. A technical quality-assurance plan that addresses all aspects of sample collection, analysis, and reporting is being prepared at the national level. This plan will be appended as needed at the pilot-project level. A National Coordinating Work Group that functions under the auspices of the Interagency Advisory Committee on Water Data and the Advisory Committee on Water Data for Public Use has been established at the national level. A local liaison committee consisting of representatives from Federal, State, and local agencies has been established to enhance communication and to ensure that the scientific information produced by the

Open-File Report

The thermal landscape of the Willamette River—Patterns and controls on stream temperature and implications for flow management and cold-water salmonids

Water temperature is a primary control on the health, diversity, abundance, and distribution of aquatic species, but thermal degradation resulting from anthropogenic influences on rivers is a challenge to threatened species worldwide. In the Willamette River Basin, northwestern Oregon, spring-run Chinook salmon ( Oncorhynchus tshawytscha ) and winter-run steelhead ( O. mykiss ) are formerly abundant cold-water-adapted species that are now protected under the Endangered Species Act. Among the challenges to the health of cold-water salmonids in the Willamette River Basin, disruptions in the seasonal patterns of stream temperature imposed by 13 large, multipurpose dams on tributaries to the Willamette River, as well as temperatures routinely in excess of regulatory limits in the Willamette River Basin, are contributing factors. To better understand controls on stream temperature, the sensitivity of stream temperature to flow augmentation as a management tool for suppressing high temperatures, and the implications for threatened salmonids, this study used a two-dimensional hydrodynamic and water-quality model (CE-QUAL-W2) to investigate spatial and temporal patterns of stream temperature in the Willamette River Basin. This study focused on the upper 160.4 river miles of the Willamette River from the confluence of the Middle Fork and Coast Fork Willamette Rivers (river mile 187.2) to Willamette Falls (river mile 26.8), three representative climate years (2011, a cool and wet year; 2015, an extremely hot and dry year; and 2016, a moderately hot and dry year), and a series of flow-augmentation scenarios. Model results show that the Willamette River upstream from Willamette Falls is divisible into four characteristic “thermal reaches” with similar thermal patterns, depending on tributary input, warming rate, and the timing of thermal response. In general, the Willamette River warms downstream during spring and summer, but patterns are complex, influenced by tributary inflows, and seasonally variable. Except in cool wet years (as illustrated by 2011), modeling suggests that adversely warm conditions for spring-run Chinook salmon are extensive from June or July through August. The thermal influence of flow augmentation from dam storage on four tributaries with U.S. Army Corps of Engineers dams varies spatially along the Willamette River, seasonally, and in magnitude, depending on a range of factors like distance from the Willamette River, the temperature of dam outflow, and the thermal template of tributary reaches controlling stream temperature adjustment to environmental heat fluxes. Modeling suggests that targeted flow management (via augmentation from dam storage) can reduce the extent and duration of thermally stressful conditions for Chinook salmon for short periods, but modeling suggests that flow augmentation is limited in its ability to fundamentally alter the “thermal landscape” (the entire range of temperature variation in a river system over space and time) of the Willamette River. While this research provides general insights into the thermal landscape of the Willamette River and its sensitivity to flow management, additional investigation into the thermal landscape of tributaries downstream from U.S. Army Corps of Engineers dams, as well as the thermal management of reservoirs, storage availability, and dam outflows, would be necessary to target specific management actions for supporting specified rearing or migration conditions for spring-run Chinook salmon and other cold-water-adapted species in the Willamette River Basin.

Oregon

Sperm cryopreservation in fish and shellfish

Initial success in sperm cryopreservation came at about the same time for aquatic species and livestock. However, in the 50-plus years since then cryopreserved sperm of livestock has grown into a billion-dollar global industry, while despite work in some 200 species with well over 200 published reports, cryopreservation of aquatic species sperm remains essentially a research activity with little commercial application. Most research has focused on large-bodied culture and sport fishes, such as salmonids, carps, and catfishes, and mollusks such as commercially important oyster and abalone species. However, only a handful of studies have addressed sperm cryopreservation in small fishes, such as zebrafish, and in endangered species. Overall, this work has yielded techniques that are being applied with varying levels of success around the world. Barriers to expanded application include a diverse and widely distributed literature base, technical problems, small sperm volumes, variable results, a general lack of access to the technology, and most importantly, the lack of standardization in practices and reporting. The benefits of cryopreservation include at least five levels of improvements for existing industries and for creation of new industries. First, cryopreservation can be used to improve existing hatchery operations by providing sperm on demand and simplifying the timing of induced spawning. Second, frozen sperm can enhance efficient use of facilities and create new opportunities in the hatchery by eliminating the need to maintain live males, potentially freeing resources for use with females and larvae. Third, valuable genetic lineages such as endangered species, research models, or improved farmed strains can be protected by storage of frozen sperm. Fourth, cryopreservation opens the door for rapid genetic improvement. Frozen sperm can be used in breeding programs to create improved lines and shape the genetic resources available for aquaculture. Finally, cryopreserved sperm of aquatic species will at some point become an entirely new industry itself. A successful industry will require integrated practices for sample collection, refrigerated storage, freezing, thawing, rules for use and disposal, transfer agreements, and database development. Indeed the development of this new industry is currently constrained by factors including the technical requirements for scaling-up to commercial operations during the transition from research, and the absence of uniform quality control practices, industry standards, marketing and price structures, and appropriate biosecurity safeguards.

Society for Reproduction and Fertility Supplement

Riverscape heterogeneity shapes population diversity for a migratory fish

Habitat patch dynamics can scale up to influence population demography and diversity with implications for resilience to environmental stochasticity. But how the spatial arrangement and size of habitat patches interact with other components of habitat heterogeneity to shape population diversity at larger spatial scales is not well understood. For riverine fishes, there is increasing evidence that tributary streams provide critical demographic support to main stem rivers. However, the extent to which main stem rivers rely on demographic contributions from tributaries, and the factors underlying this dependence, have not been assessed. Here, we used genetic stock identification to evaluate the effect of tributaries on population diversity of Yellowstone cutthroat trout ( Oncorhynchus virginalis bouvieri ) occupying the main stem Snake River, Wyoming, USA. We found that the main stem relied almost entirely on tributaries for demographic support, but main stem composition varied spatially among river sections. Distance between habitat patches, catchment area, and groundwater availability acted in concert to determine the contribution of specific tributaries to the main stem, but contributions were ultimately modulated by habitat connectivity. We also found evidence for multi-scale spatial structure in tributary contributions, providing insight into untested drivers of main stem river population diversity. Our results demonstrate how spatially discrete and distributed riverscape attributes influence population diversity at broader spatial scales, illustrating how ecosystem resilience emerges from the dynamic, two-way exchange of individuals and energy across habitat networks. Management plans for large rivers that address the ecological contributions of tributaries may be needed to achieve optimal outcomes. Similarly, conservation strategies that exclusively focus on headwater streams may fail to capture the broader habitat requirements necessary to maintain robust cold-water fish populations and associated recreational fisheries, particularly under global environmental change.

Wyoming

Landscape-scale determinants of native and nonnative Great Plains fish distributions

Aim Landscape-scale factors may have differential effects on the distribution of native and non-native fishes and may help explain invasion success and species declines. Location Great Plains, Wyoming, USA Methods We used hierarchical Bayesian mixture models and constrained ordination techniques to evaluate associations between landscape-scale factors on native and non-native fish species richness, reproductive guilds and individual species distributions. Results Predicted responses between landscape-scale factors and native and non-native fish species richness were similar, except non-native fish species richness that was positively associated with density of oil and gas wells. Non-native fish species richness was also positively associated with native fish species richness. Spawning guild composition differed between native and non-native fishes. Canonical correspondence analysis revealed that the most abundant non-native and only a few native species were positively associated with oil and gas wells. Main conclusions The similar relationships between native and non-native fish species richness are likely evidence that they share similar ecological rules, which supports that non-native species become naturalized and they may be affected by the same environmental factors that determine distribution of native species.

Diversity and Distributions

Modeling ecological minimum requirements for distribution of greater sage-grouse leks: implications for population connectivity across their western range, U.S.A.

Greater sage-grouse Centrocercus urophasianus (Bonaparte) currently occupy approximately half of their historical distribution across western North America. Sage-grouse are a candidate for endangered species listing due to habitat and population fragmentation coupled with inadequate regulation to control development in critical areas. Conservation planning would benefit from accurate maps delineating required habitats and movement corridors. However, developing a species distribution model that incorporates the diversity of habitats used by sage-grouse across their widespread distribution has statistical and logistical challenges. We first identified the ecological minimums limiting sage-grouse, mapped similarity to the multivariate set of minimums, and delineated connectivity across a 920,000 km 2 region. We partitioned a Mahalanobis D 2 model of habitat use into k separate additive components each representing independent combinations of species–habitat relationships to identify the ecological minimums required by sage-grouse. We constructed the model from abiotic, land cover, and anthropogenic variables measured at leks (breeding) and surrounding areas within 5 km. We evaluated model partitions using a random subset of leks and historic locations and selected D 2 (k = 10) for mapping a habitat similarity index (HSI). Finally, we delineated connectivity by converting the mapped HSI to a resistance surface. Sage-grouse required sagebrush-dominated landscapes containing minimal levels of human land use. Sage-grouse used relatively arid regions characterized by shallow slopes, even terrain, and low amounts of forest, grassland, and agriculture in the surrounding landscape. Most populations were interconnected although several outlying populations were isolated because of distance or lack of habitat corridors for exchange. Land management agencies currently are revising land-use plans and designating critical habitat to conserve sage-grouse and avoid endangered species listing. Our results identifying attributes important for delineating habitats or modeling connectivity will facilitate conservation and management of landscapes important for supporting current and future sage-grouse populations.

Ecology and Evolution

The rise of an apex predator following deglaciation

Aim Sea otters ( Enhydra lutris ) are an apex predator of the nearshore marine community and nearly went extinct at the turn of the 20th century. Reintroductions and legal protection allowed sea otters to re‐colonize much of their former range. Our objective was to chronicle the colonization of this apex predator in Glacier Bay, Alaska, to help understand the mechanisms that governed their successful colonization. Location Glacier Bay is a tidewater glacier fjord in southeastern Alaska that was entirely covered by glaciers in the mid‐18th century. Since then, it has endured the fastest tidewater glacier retreat in recorded history. Methods We collected and analysed several data sets, spanning 20 years, to document the spatio‐temporal dynamics of an apex predator expanding into an area where they were formerly absent. We used novel quantitative tools to model the occupancy, abundance and colonization dynamics of sea otters, while accounting for uncertainty in the data collection process, the ecological process and model parameters. Results Twenty years after sea otters were first observed colonizing Glacier Bay, they became one of the most abundant and widely distributed marine mammal. The population grew exponentially at a rate of 20% per year. They colonized Glacier Bay at a maximum rate of 6 km per year, with faster colonization rates occurring early in the colonization process. During colonization, sea otters selected shallow areas, close to shore, with a steep bottom slope, and a relatively simple shoreline complexity index. Main conclusions The growth and expansion of sea otters in Glacier Bay demonstrate how legal protection and translocation of apex predators can facilitate their successful establishment into a community in which they were formerly absent. The success of sea otters was, in part, a consequence of habitat that was left largely unperturbed by humans for the past 250 years. Further, sea otters and other marine predators, whose distribution is limited by ice, have the potential to expand in distribution and abundance, reshaping future marine communities in the wake of deglaciation and global loss of sea ice.

Alaska

Patterns of species richness hotspots and estimates of their protection are sensitive to spatial resolution

Aim Species richness is a measure of biodiversity often used in spatial conservation assessments and mapped by summing species distribution maps. Commission errors inherent those maps influence richness patterns and conservation assessments. We sought to further the understanding of the sensitivity of hotspot delineation methods and conservation assessments to commission errors, and choice of threshold for hotspot delineation. Location United States. Methods We created range maps and 30‐m and 1‐km resolution habitat maps for terrestrial vertebrates in the United States and generated species richness maps with each dataset. With the richness maps and the GAP Protected Areas Dataset, we created species richness hotspot maps and calculated the proportion of hotspots within protected areas; calculating protection under a range of thresholds for defining hotspots. Our method allowed us to identify the influence of commission errors by comparing hotspot maps. Results Commission errors from coarse spatial grain data and lack of porosity in the range data inflated richness estimates and altered their spatial patterns. Coincidence of hotspots from different data types was low. The 30‐m hotspots were spatially dispersed, and some were very long distances from the hotspots mapped with coarser data. Estimates of protection were low for each of the taxa. The relationship between estimates of hotspot protection and threshold choice was nonlinear and inconsistent among data types (habitat and range) and grain size (30‐m and 1‐km). Main conclusions Coarse mapping methods and grain sizes can introduce commission errors into species distribution data that could result in misidentifications of the regions where hotspots occur and affect estimates of hotspot protection. Hotspot conservation assessments are also sensitive to choice of threshold for hotspot delineation. There is value in developing species distribution maps with high resolution and low rates of commission error for conservation assessments.

Diversity and Distributions

Predictive multi-scale occupancy models at range-wide extents: Effects of habitat and human disturbance on distributions of wetland birds

Aim Predicting distributions is fundamental to ecology, yet hindered by spatially restricted sampling, scale-dependent relationships and detection error associated with field surveys. Predictive species distribution models (SDMs) are nonetheless vital for conservation of many species. We developed a framework for building predictive SDMs with multi-scale data and used it to develop range-wide breeding-season SDMs for 14 marsh bird species of concern. Location USA. Methods We built SDMs using data from range-wide surveys conducted over 14 years, and habitat and disturbance covariates measured at multiple spatial scales. We built hierarchical occupancy models that included heterogeneity in detectability during sampling, and used Bayesian model selection to regulate model complexity (covariates and scales) based explicitly on spatial predictive abilities. We thus integrated model selection for optimizing out-of-sample prediction, range-wide sampling over broad conditions, multi-scale analyses and scale optimization, and species-specific detectability for a suite of wide-ranging species. Results Distributions of marsh birds were affected by local wetland conditions, but also by agricultural, urban and hydrologic disturbances operating from local scales (100–500 m) to the watershed level. Variables measuring human disturbances improved prediction for most species, and every species was affected by attributes at >1 scale. Five species showed evidence for continental-scale range contraction during the study. Main conclusions We demonstrate how hierarchical occupancy models can be optimized for prediction across a species' range at the extent of a continent while also accounting for imperfect detection, and thus describe a generalizable approach that can be used for any species. We provide the first data-driven, empirical SDMs built at the range-wide extent for most of our 14 study species and demonstrate that previous studies focused on local distributions and the effects of fine-scale wetland vegetation missed important broadscale drivers of occupancy for marsh birds.

Diversity and Distributions