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Thin-layer sediment addition to an existing salt marsh to combat sea-level rise and improve endangered species habitat in California, USA

Current tidal marsh elevations and their accretion rates are important predictors of vulnerability to sea-level rise. When tidal marshes are at risk, adaptation measures, such as sediment addition to increase elevations, can be implemented to prevent degradation and loss. In 2016, wildlife managers prescribed a thin-layer sediment addition of locally sourced dredged material from Anaheim Bay to mitigate plausible future impacts of sea-level rise to Seal Beach National Wildlife Refuge, an urbanized, subsiding, sediment limited, and low elevation tidal marsh in southern California, USA. In this study, our objectives were to determine how suspended sediment concentrations (SSC) and fluxes associated with sediment application affected an adjacent eelgrass bed and a more distant deep tidal channel at different time periods throughout the project and how sediment application affected tidal marsh elevation. Due to the sediment addition the tidal marsh elevation increased by 25.4 cm across 3.06 ha. Mean SSC levels at the eelgrass site increased from 4 mg/L during pre-augmentation to 16 mg/L during sediment application. SSC levels also increased up to 40 mg/L with the installation of sediment barriers. Winter storms during construction also had an influence on SSC, with an extreme storm event increasing SSC from 9 mg/L pre-storm levels to 16 mg/L during the storm. However, SSC at the deep site remained constant during all time periods with a mean of 7 mg/L. Our results show that little to zero sediment was exported out of Anaheim Bay during the study period, illustrating that any impacts were localized to the application area. These findings suggest that although long-range export of applied dredge material is unlikely from future thin-layer sediment application at this site, adjacent habitats could experience impacts if erosion control measures are not implemented effectively. Future projects will benefit from similar pre- and post-addition monitoring to establish the long-term efficacy of sea-level rise adaptation measures.

California

Status review of the Marbled Murrelet (Brachyramphus marmoratus) in Alaska and British Columbia

The Marbled Murrelet ( Brachyramphus marmoratus ) is a small, diving seabird inhabiting inshore waters of the Northeastern Pacific Ocean. This species feeds on small, schooling fishes and zooplankton, and nests primarily on the moss-covered branches of large, old-growth conifers, and also, in some parts of its range, on the ground. We reviewed existing information on this species to evaluate its current status in the northern part of its range—Alaska (U.S.) and British Columbia (Canada). Within the southern part of its range (Washington, Oregon, and California, U.S.), the Marbled Murrelet was listed as a threatened species under the Endangered Species Act (ESA) in 1993, and the U.S. Fish and Wildlife Service (USFWS) needed information on the species throughout its range for ESA deliberations. We compiled published information on the conservation status, population biology, foraging ecology, population genetics, population status and trends, demography, marine and nesting habitat characteristics, threats, and ongoing conservation efforts for Marbled Murrelets in Alaska and British Columbia. We conducted a new genetic study using samples from a segment of the range that had not been included in previous studies (Washington, Oregon) and additional nuclear intron and microsatellite markers. We also analyzed available at-sea survey data from several locations for trend. To understand the reasonableness of the empirical trend data, we developed demographic models incorporating stochasticity to discern what population trends were possible by chance. The genetic studies substantially confirmed previous findings on population structure in the Marbled Murrelet. Our present work finds three populations: (1) one comprising birds in the central and western Aleutian Islands; (2) one comprising birds in central California; and (3) one comprising birds within the center of the range from the eastern Aleutians to northern California. Our knowledge of genetic structure within this central population is limited and it requires additional study. Compiling available abundance information, we estimated that in the recent past, Marbled Murrelets in Alaska numbered on the order of 1 million birds. We were unable to generate a similar estimate for historical population size in British Columbia. Using trend information from at-sea surveys spanning a wide geographic range in Alaska, murrelet numbers declined significantly at five of eight trend sites at annual rates of -5.4 to -12.7 percent since the early 1990s. Applying these rates of decline to the historical population estimate, the current murrelet population in Alaska is projected to be on the order of 270,000 birds. This represents an overall population decline of about 70 percent during the past 25 years. In British Columbia, available trend data indicate that murrelet populations there have experienced similar declines. We updated a recent (2002) population estimate for British Columbia, concluding that there are now between 54,000 and 92,000 murrelets in British Columbia. The rates of decline we observed are within, but at the high end of, a range of rates expected by chance. Given that declines were estimated for sites over essentially the entire northern range of the species, there is cause for concern about the species’ status. In their marine habitats, Marbled Murrelets overlap with salmon (Oncorhynchus sp.) gillnetting operations in British Columbia and in Alaska (especially in Prince William Sound and Southeast Alaska), and annual bycatch mortality is likely in the low thousands per year, although bycatch rates are difficult to measure. The species’ inshore distribution coincides with high levels of vessel traffic and makes them especially vulnerable to both chronic oil pollution and to catastrophic spills (e.g., the 1989 Exxon Valdez oil spill [EVOS] in south-central Alaska, which is estimated to have killed 12,000 to 15,000 murrelets). In their forested nesting habitats, Marbled Murrelets have lost about 15 percent of their suitable nesting habitat in Southeast Alaska, and 33 to 49 percent in British Columbia, from industrial-scale logging within the past half century. Increased predation also may be a threat to murrelet populations, related to fragmentation and edge effects from logging and development, and recent population increases observed for some important murrelet predators, including Bald Eagles ( Haliaeetus leucocephalus ), Common Ravens ( Corvus corax ), and Steller’s Jays ( Cyanocitta stelleri ). Nesting habitat losses cannot explain the declines observed in areas where industrial logging has not occurred on a large scale (e.g., Prince William Sound) or at all (Glacier Bay). The apparent change in population size and rates of decline reported for the Marbled Murrelet are large, and we therefore considered alternative explanations and precedents for changes of similar magnitude in other marine wildlife populations in the Northeastern Pacific Ocean. The declines are likely real, and related to combined and cumulative effects from climate-related changes in the marine ecosystem (most likely the 1977 regime shift) and human activities (logging, gillnet bycatch, oil pollution). Much uncertainty about the decline could be alleviated by continuing to repeat boat surveys in Prince William Sound and lower Cook Inlet, and by repeating the boat survey of Southeast Alaska that was conducted in 1994. This survey used a statistically sound design and covered the region that has been and likely remains the center of the species’ abundance. Important questions remain to be addressed about methods for measuring population status and change, adult mortality (major sources, density dependence, seasonal concordance), and the movements of wintering populations.

Alaska, British Columbia

Evaluation of alternative groundwater-management strategies for the Bureau of Reclamation Klamath Project, Oregon and California

The water resources of the upper Klamath Basin, in southern Oregon and northern California, are managed to achieve various complex and interconnected purposes. Since 2001, irrigators in the Bureau of Reclamation Klamath Irrigation Project (Project) have been required to limit surface-water diversions to protect habitat for endangered freshwater and anadromous fishes. The reductions in irrigation diversions have led to an increased demand for groundwater by Project irrigators, particularly in drought years. The potential effects of sustained pumping on groundwater and surface-water resources have caused concern among Federal and state agencies, Indian tribes, wildlife groups, and groundwater users. To aid in the development of a viable groundwater-management strategy for the Project, the U.S. Geological Survey, in collaboration with the Klamath Water and Power Agency and the Oregon Water Resources Department, developed a groundwater-management model that links groundwater simulation with techniques of constrained optimization. The overall goal of the groundwater-management model is to determine the patterns of groundwater pumping that, to the extent possible, meet the supplemental groundwater demands of the Project. To ensure that groundwater development does not adversely affect groundwater and surface-water resources, the groundwater-management model includes constraints to (1) limit the effects of groundwater withdrawal on groundwater discharge to streams and lakes that support critical habitat for fish listed under the Endangered Species Act, (2) ensure that drawdowns do not exceed limits allowed by Oregon water law, and (3) ensure that groundwater withdrawal does not adversely affect agricultural drain flows that supply a substantial portion of water for irrigators and wildlife refuges in downslope areas of the Project. Groundwater-management alternatives were tested and designed within the framework of the Klamath Basin Restoration Agreement (currently [2013] awaiting authorizing Federal legislation), which would establish a permanent limit on the amount of surface water that can be diverted annually to the Project. Groundwater-management scenarios were evaluated for the period 1970•2004; supplemental groundwater demand by the Project was estimated as the part of irrigation demand that would not have been satisfied by the surface-water diversion allowed under the Klamath Basin Restoration Agreement. Over the 35-year management period, 22 years have supplemental groundwater demand, which ranges from a few thousand acre-feet (acre-ft) to about 100,000 acre-ft in the driest years. The results of the groundwater-management model indicate that supplemental groundwater pumping by the Project can be managed to avoid adverse effects to groundwater discharge that supports critical aquatic habitat. The existing configuration of wells in the Project would be able to meet groundwater-pumping goals in 14 of the 22 years with supplemental groundwater demand; however, substantial irrigation shortages can be expected during drought periods when the demand for supplemental groundwater is highest. The maximum irrigation-season withdrawal calculated by the groundwater-management model is about 60,000 acre-ft, the average withdrawal in drought years is about 54,000 acre-ft, and the amount of unmet groundwater demand reaches a maximum of about 45,000 acre-ft. A comparison of optimized groundwater withdrawals by geographic region shows that the highest annual withdrawals are associated with wells in the Tule Lake and Klamath Valley regions of the Project. The patterns of groundwater withdrawal also show that a substantial amount of the available pumping capacity is unused due to the restrictions imposed by drawdown constraints. Subsequent model applications were used to evaluate the sensitivity of optimization results to various factors. A sensitivity analysis quantified the changes in optimized groundwater withdrawals that result from changes in drawdown-constraint limits. The analysis showed the potential for substantial increases in withdrawals of groundwater with less restrictive drawdown limits at drawdown-control sites in the California part of the model. Systematic variation of the drains-constraint limit yielded a trade-off curve between optimized groundwater withdrawals and the allowable reduction in groundwater discharge to the Project drain system. Additional model applications were used to assess the value of increasing the pumping capacity of the network of wells serving the Project, and the relation between reduced off-Project groundwater pumping and increased pumping by Project irrigators.

California;Oregon

Modeling Least Bell’s Vireo habitat suitability in current and historic ranges in California

We developed a habitat suitability model for the federally endangered Least Bell’s Vireo ( Vireo bellii pusillus ) across its current and historic range in California. The vireo disappeared from most of its range by the 1980s, remaining only in southern California and northern Baja California, Mexico. This decline was due to habitat loss and introduction of brood parasitic brown-headed cowbirds ( Molothrus ater ) into California in the late 1800s. Habitat protection and management since the mid-1980s increased southern California vireo populations with small numbers of birds recently expanding back into the historic range. The vireo habitat model will help meet the U.S. Fish and Wildlife Service recovery objectives by distinguishing specific areas to survey for new occurrences; characterizing important vireo-habitat relationships; and identifying areas for habitat management. We constructed models based on the vireo’s current range to predict suitable habitat in the historic range, which differs substantially in environmental conditions. We used the partitioned Mahalanobis D 2 modeling technique designed to predict habitat suitability in areas not included in a sample of species locations and under novel conditions. We constructed alternative models with different combinations of environmental variables hypothesized to be important components of vireo habitat. We selected a set of best performing models to predict suitable habitat for a riparian vegetation grid buffered 500 meters across California. Most models for southern California did not predict suitable habitat in the historic range. The top performing model has an area under the curve value of 0.93. It is a simple model and discriminated among riparian habitats, with only 6 percent predicted as suitable. On average, suitable vireo habitat had more than 60-percent riparian vegetation and flat land at the 150-meter scale, little-to-no slope, and was within 130 meters of water.

California

Status of the Island Night Lizard and Two Non-Native Lizards on Outlying Landing Field San Nicolas Island, California

More than 900 individually marked island night lizards (Xantusia riversiana) were captured on San Nicolas Island, California, between 1984 and 2007 as part of an ongoing study to monitor the status of this threatened species. Our data suggest that at least a few lizards are probably more than 20 years old, and one lizard would be 31.5 years old if it grew at an average rate for the population. Ages of 20 and 30 years seem reasonable given the remarkably slow growth during capture intervals of more than a decade for five of the lizards which we estimated to be 20 or more years old. Like other lizards, island night lizard growth rates vary by size, with larger lizards growing more slowly. In general, growth rates were somewhat greater on San Nicolas Island (compared with Santa Barbara Island), and this increase was sustained through all of the intermediate size classes. The higher growth rate may account for the somewhat larger lizards present on San Nicolas Island, although we cannot discount the possibility that night lizards on San Nicolas are merely living longer. The high percentage of small lizards in the Eucalyptus habitat might seem to reflect a healthy population in that habitat, but the high proportion of small lizards appears to be caused by good reproduction in the 1900s and substantially poorer reproduction in subsequent years. The Eucalyptus habitat has dried quite a bit in recent years. Night lizards in the Haplopappus/Grassland habitat have shown an increase in the proportion of larger lizards since 2000. There has also been an increase in the proportion of large lizards in the Rock Cobble habitat at Redeye Beach. However, there are has been some change in habitat with more elephant seals occupying the same area just above the high tide as do the night lizards. Southern alligator lizards and side-blotched lizards are both non-native on San Nicolas Island. Neither lizard causes obvious harm to island night lizards, and management time and effort should be directed toward much more pressing problems, such as general habitat restoration, erosion control, and the removal of feral cats. The island night lizard (Xantusia riversiana) is endemic to three of the California Channel Islands: Nicolas, San Clemente, and Santa Barbara Islands. Due to its restricted range and apparently small population levels, both the U.S. Fish and Wildlife Service and the California Department of Fish and Game have listed the island night lizard as a threatened species. Our study was conducted on San Nicolas Island, which lies offshore 120 km southwest of Los Angeles, California. The island is managed by the U.S. Navy who refers to the island as Outlying Landing Field San Nicolas Island. The Navy maintains radar, telemetry, and communications equipment on San Nicolas Island to support its mission of testing and evaluating weapons systems. The Navy has dual requirements for ensuring military readiness and sustainability while complying with the Federal Endangered Species Act. A comprehensive understanding of the status and stability of the species on San Nicolas Island is essential for effective island management and may aid in the eventual delisting of the species. Previous work on the San Nicolas Island (Fellers and others, 1998) demonstrated that island night lizards were distributed over the eastern half of San Nicolas Island where there is suitable shrubby habitat. On the eastern half of the island, they occur primarily in or near cactus/sage scrub habitats on the north beach terrace, in scattered patches of scrub on the central mesa, and in boulder and cactus habitats on the southern escarpment of the island. Fellers and others (1998) evaluated data from 1984-85 and 1992-95 and estimated that there were 15,300 island night lizards present on San Nicolas Island. There are two non-native lizards on San Nicolas Island, the side-blotch lizard (Uta stansburiana) and the southern alligator lizard (Elgaria multicarinata). Both of the

Open-File Report

Developing an effective Agassiz's Desert Tortoise monitoring program: Final report to the Coachella Valley Conservation Commission

Agassiz’s desert tortoise (Gopherus agassizii) is a conservation-reliant species with populations north and west of the Colorado River protected as threatened under the Endangered Species Act (Averill-Murray et al. 2012). Since it was listed under this category in 1990, a great deal has been learned about the natural history of the species, and it is now one of the best-studied turtles in the United States (Lovich and Ennen 2013). However, the accumulated body of scientific data available for the species has not yet been translated into recovery or delisting of the species. Successful conservation of any species requires knowledge of their natural history and how vital rates affect their ability to maintain stable populations in the face of natural and anthropogenic stresses. Agassiz’s desert tortoises occur from southwestern Utah to near the Mexican border in California – a distance of over 450 km – but population densities vary greatly across this immense landscape (U.S. Fish and Wildlife Service 2015). Tortoises occur in the Sonoran Desert of California, including the eastern and western ends of the Coachella Valley, where it is one of 27 species covered under the Coachella Valley Multiple Species Habitat Conservation Plan and Natural Community Conservation Plan (CVMSHCP/NCCP). The southern portion of Joshua Tree National Park (JTNP) lies within this 1.1 million acre planning area, and was predicted to be an area of low-density tortoise populations using habitat suitability modeling (Barrows 2011). JTNP is near the southern distributional limit of G. agassizii, yet very little has been published regarding the ecology of tortoises in the Sonoran Desert of California. Reproductive output is an important gross measure of the ability of a population to persist. When integrated with data on fertility and survivorship, this information forms a foundation for assessing population status and formulating effective management strategies (e.g., Congdon et al. 1993, 1994), especially for imperiled species. One aspect of the biology of G. agassizii that has been particularly well-studied is reproductive output. However, most of what we know about this topic comes from research in the Mojave Desert portion of the species’ range (Ernst and Lovich 2009). Comparatively little has been published on the reproductive ecology of populations living in the Sonoran Desert ecosystem of California. Publications by Lovich et al. (1999, 2011, 2012, 2014, 2015) constitute the main body of literature on desert tortoise reproductive ecology in the Sonoran Desert of California, with one study population located at the western end of the CVMSHCP/NCCP area. Collecting data on Agassiz’s desert tortoise ecology in the Sonoran Desert ecosystem is important due to significant differences between the two adjacent desert ecosystems, especially the timing and amounts of annual precipitation, and their potential effects on reproductive output (e.g., Lovich et al. 5 2015). There are also differences in the vulnerability of tortoises to the effects of a warming, drying climate between the two deserts (Barrows 2011; Zylstra et al. 2012). The overall goal of this study was to collect data on demography, reproductive output, and genetic affinities at a study site in the Sonoran Desert portion of JTNP in the eastern end of the CVMSHCP/NCCP area. Specific objectives included: 1) Collect data to establish baselines on tortoise populations and/or their habitat suitability in core habitat within the CVNCCP area, including biotic and abiotic variables affecting persistence of tortoise populations; 2) Compare and contrast with data collected on desert tortoises at USGS/BLM study site near Palm Springs over 16 years; 3) Support long-term modeling efforts needed to determine tortoise population viability; 4) Refine modeled relationships with identified threats such as fire, invasive species and climate change; and 5) Prioritize adaptive management needs for the desert tortoise in and beyond the CVNCCP area. The data from this study will aid in determining baseline estimates of the desert tortoise population size within the planning area as well as establish a marked population of Agassiz’s desert tortoises for future monitoring. Data will be integrated with habitat modeling in order to refine model output. Genetic data will be collected on both the north and south sides of Interstate 10 to determine the potential effects of habitat fragmentation and genetic mixing. Analyses are ongoing and results beyond those presented in this report will be published in peer-reviewed scientific journals following inclusion of additional data collected on the south side of Shavers Valley in 2017-2018.

Report

Persistence of historical population structure in an endangered species despite near-complete biome conversion in California's San Joaquin Desert

Genomic responses to habitat conversion can be rapid, providing wildlife managers with time-limited opportunities to enact recovery efforts that use population connectivity information that reflects predisturbance landscapes. Despite near-complete biome conversion, such opportunities may still exist for the endemic fauna and flora of California's San Joaquin Desert, but comprehensive genetic data sets are lacking for nearly all species in the region. To fill this knowledge gap, we studied the rangewide population structure of the endangered blunt-nosed leopard lizard Gambelia sila , a San Joaquin Desert endemic, using restriction site-associated DNA (RAD), microsatellite and mtDNA data to test whether admixture patterns and estimates of effective migration surfaces (EEMS) can identify land areas with high population connectivity prior to the conversion of native xeric habitats. Clustering and phylogenetic analyses indicate a recent shared history between numerous isolated populations and EEMS reveals latent signals of corridors and barriers to gene flow over areas now replaced by agriculture and urbanization. Conflicting histories between the mtDNA and nuclear genomes are consistent with hybridization with the sister species G. wislizenii , raising important questions about where legal protection should end at the southern range limit of G. sila . Comparative analysis of different data sets also adds to a growing list of advantages in using RAD loci for genetic studies of rare species. We demonstrate how the results of this work can serve as an evolutionary guidance tool for managing endemic, arid-adapted taxa in one of the world's most compromised landscapes.

California

Population and habitat analyses for greater sage-grouse (Centrocercus urophasianus) in the bi-state distinct population segment—2018 update

Executive Summary The Bi-State Distinct Population Segment (Bi-State DPS) of greater sage-grouse ( Centrocercus urophasianus , hereinafter “sage-grouse”) represents a genetically distinct and geographically isolated population that straddles the border between Nevada and California. The primary threat to these sage-grouse populations is the expansion of single-leaf pinyon ( Pinus monophylla ) and Utah juniper ( Juniperus osteosperma ) into sagebrush ecosystems, which fragments and reduces population connectivity and survival. Other important threats include low water availability during brood-rearing, particularly during drought, and increased predation by common ravens ( Corvus corax ), a generalist predator often associated with anthropogenic resource subsidies. Although the Bi-State DPS occurs at high elevations relative to sage-grouse range-wide, changes in historical wildfire cycles and the conversion of native shrubs to invasive annual grasslands still threaten these populations. The Bi-State DPS has undergone multiple federal status assessments and associated litigation. For example, in October of 2013, the Bi-State DPS was proposed for listing as threatened under the Endangered Species Act of 1973 by the U.S. Fish and Wildlife Service (USFWS), then withdrawn in April 2015. The withdrawal decision was challenged, and in May 2018, a Federal district court ordered the withdrawal decision to be vacated, and USFWS was required to re-open the October 2013 listing evaluation. In response, the U.S. Geological Survey (USGS), with State and Federal collaborators, embarked on a multipronged analysis to provide current and best available science regarding population status of sage-grouse within the Bi-State DPS. Using data from a long-term monitoring program, we carried out four analytical study objectives, and here, we provide preliminary results of these analyses. First, we used integrated population modeling (IPM) to predict annual population abundance and annual finite rate of population change for the Bi-State DPS, as a whole, and for each subpopulation between 1995 and 2018. Because sage-grouse exhibit population cycles (periodic increases and decreases in abundance across approximately 6- to 10-year wavelengths), we estimated trends across three nested temporal scales that represent one (11 years), two (18 years), and three (24 years) complete population cycles. These estimates of relatively long-term averaged population change account for temporal (that is, interannual) variation. Our model predicted population abundance for the Bi-State DPS during 2018 at 3,305 individuals (2,247–4,683), with the majority occupying Bodie Hills and Long Valley. The model also predicted cyclic dynamics in abundance through time with evidence of 24-year population growth and slight trends of decline over the past 18 years. Specifically, across the Bi-State DPS as a whole, we estimated annual average at 0.99, 0.99, and 1.02 over the one, two, and three population cycles, which equated to a 10.5 percent, 16.6 percent decrease, and 60.0 percent increase in abundance over the 11-, 18-, and 24-year cycles. Estimated abundance in 2018 had not reached numbers lower than those predicted during 1995. However, we found spatial variation in population trends across the three cycles. Bodie Hills subpopulation comprised the greatest (1,521) and exhibited average annual greater than 1.0 across all periods resulting in average annual increases of 7 percent. This relatively large subpopulation has grown 5 times larger than what was predicted in 1995 while experiencing cyclical dynamics within that period. Conversely, other smaller subpopulations within the Bi-State DPS exhibited average annual equal to or less than 1.0 resulting in estimated 10-year risks of extirpation ranging from 2.0 to 76.1 percent. In general, evidence of decline among smaller subpopulations was greatest for the most recent period (2008–18) compared to a period that encompassed three full population cycles (24-year). This difference coincides with an intense period of drought that began in 2012. For comparative purposes as part of this first objective, we conducted a similar analysis for populations of sage-grouse within Nevada and California but outside the Bi-State DPS. We developed a region-wide and distance-weighted IPM using lek count from Nevada Department of Wildlife (NDOW) and California Department of Fish and Wildlife (CDFW) databases and with telemetry data collected by USGS across 12 sage-grouse subpopulations. Our models predicted similar patterns in population cycling outside the Bi-State DPS but with much stronger evidence of long-term declines across 24 years. Specifically, median averaged across each year of the 11-, 18-, and 24-year periods resulted in average annual values of 0.94, 0.97, and 0.99, respectively. These values equate to 41.0 percent, 38.5 percent, and 21.3 percent declines over the corresponding periods. Second, we used lek count data in a state-space modeling framework to compare trends in population abundance across different spatial scales (that is, leks versus Bi-State DPS). This hierarchical framework allowed us to disentangle declines associated with climate conditions as opposed to other local level factors that might signal the need for management intervention. Specifically, we identified 7 leks that were both declining and recently decoupled from larger spatial scale trends, typically governed by climatic conditions (referred to as soft or hard signals). The goal of this analysis was to provide an early warning system that might have implications for conservation actions at local scales. Third, we developed phenological (spring, summer–fall, and winter) and reproductive life stage (nesting, early brood-rearing, and late-brood rearing) based resource selection functions using various environmental covariates. We report rankings of variable importance for each season and life stage, developed maps of habitat selection indices (HSI), binned categories representing low, moderate, and high classes of quality (where any category greater than or equal to low indicated selected habitat) for each phenological season and life stage, and produced composite maps by selected phenological and reproductive stage to estimate annual habitat. Fourth, we used for each lek within the Bi-State DPS to carry out a spatial analysis that quantified substantial changes in the distribution of occupied habitat across long- (24-year) and short- (11-year) term periods. Owing to differences among available datasets, the long-term analysis primarily reflected spatial shifts among subpopulations comprising the majority of the Bi-State DPS (that is, Bodie Hills and Long Valley) while the short-term analysis also quantified changes among subpopulations along the periphery. Over long and short-term periods, the overall distribution of occupied habitat (as measured by 99 percent utilization distributions intersecting any quantified habitat) was reduced by 20,573 ha and 55,492 ha, respectively. Occupied core areas (as measured by 50 percent utilization distributions intersecting any quantified habitat) over long-term periods were solely located in Bodie Hills and Long Valley. Although nearly all subpopulations experienced contractions in occupied overall and core distribution, Bodie Hills experienced spatial expansion that occurred with concomitant spatial contraction at Long Valley over both periods. Subpopulations at the northern (Pine Nuts), central (Sagehen) and southern (White-Mountains) extents of the Bi-State DPS also experienced spatial contraction over the short-term period. These findings, coupled with those of population trends, indicate long-term patterns in redistribution of sage-grouse from Long Valley and peripheral subpopulations to Bodie Hills. That is, sage-grouse subpopulations at the periphery are declining while the largest population at the core is increasing, which could have meaningful impacts on overall metapopulation persistence. We provide evidence for loss of occupied habitat (reduced distribution) given local extirpation of subpopulations. Fifth, we calculated percentages of selected phenological, life stage, and annual habitat that each subpopulation contributed to the Bi-State DPS. We then intersected these maps with a composite estimate of occupied habitat from the fourth objective and calculated percentages of selected habitat likely occupied by sage-grouse that each subpopulation contributed to the Bi-State DPS. These values provide evidence for loss of occupied habitat and subsequent reductions in spatial distribution given reductions in abundance and, in some cases, extirpation of leks within subpopulations. Lastly, we carried out an initial in-depth analysis of selection for irrigated pastures and wet meadows during the brood-rearing stage for the Long Valley subpopulation. We chose this subpopulation because it represents a population core, representing 26.5 percent of total sage-grouse within the Bi-State DPS, and has exhibited long-term declines in abundance and distribution. This subpopulation is highly sensitive to precipitation and other factors that influence water availability. Models predicted higher use of the interior portions of irrigated pastures and wet meadows during late brood-rearing period, which represented a potentially risky use of habitat that was exacerbated during periods of low moisture (for example, drought, reduced water delivery, or both). Sage-grouse typically used edges of riparian areas and pastures, largely because the interior of these mesic areas consisted of considerably less overhead concealment cover (for example, shrubs) that likely resulted in a higher risk of mortality. We found that a lack of water delivery to pastures in the form of overwinter precipitation or diversion ditches increased the movements of sage-grouse to the interior of pastures. Although further investigation of water delivery impacts on chick survival are warrented, our initial findings regarding resource selection may explain recent declines in population growth at Long Valley.

California, Nevada