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The role of driving factors in historical and projected carbon dynamics of upland ecosystems in Alaska

It is important to understand how upland ecosystems of Alaska, which are estimated to occupy 84% of the state (i.e., 1,237,774 km 2 ), are influencing and will influence state‐wide carbon (C) dynamics in the face of ongoing climate change. We coupled fire disturbance and biogeochemical models to assess the relative effects of changing atmospheric carbon dioxide (CO 2 ), climate, logging and fire regimes on the historical and future C balance of upland ecosystems for the four main Landscape Conservation Cooperatives (LCCs) of Alaska. At the end of the historical period (1950–2009) of our analysis, we estimate that upland ecosystems of Alaska store ~50 Pg C (with ~90% of the C in soils), and gained 3.26 Tg C/yr. Three of the LCCs had gains in total ecosystem C storage, while the Northwest Boreal LCC lost C (−6.01 Tg C/yr) because of increases in fire activity. Carbon exports from logging affected only the North Pacific LCC and represented less than 1% of the state's net primary production (NPP). The analysis for the future time period (2010–2099) consisted of six simulations driven by climate outputs from two climate models for three emission scenarios. Across the climate scenarios, total ecosystem C storage increased between 19.5 and 66.3 Tg C/yr, which represents 3.4% to 11.7% increase in Alaska upland's storage. We conducted additional simulations to attribute these responses to environmental changes. This analysis showed that atmospheric CO 2 fertilization was the main driver of ecosystem C balance. By comparing future simulations with constant and with increasing atmospheric CO 2 , we estimated that the sensitivity of NPP was 4.8% per 100 ppmv, but NPP becomes less sensitive to CO 2 increase throughout the 21st century. Overall, our analyses suggest that the decreasing CO 2 sensitivity of NPP and the increasing sensitivity of heterotrophic respiration to air temperature, in addition to the increase in C loss from wildfires weakens the C sink from upland ecosystems of Alaska and will ultimately lead to a source of CO 2 to the atmosphere beyond 2100. Therefore, we conclude that the increasing regional C sink we estimate for the 21st century will most likely be transitional.

Alaska↗

Inter-population differences in salinity tolerance of adult wild Sacramento splittail: osmoregulatory and metabolic responses to salinity

The Sacramento splittail ( Pogonichthys macrolepidotus ) is composed of two genetically distinct populations endemic to the San Francisco Estuary (SFE). The allopatric upstream spawning habitat of the Central Valley (CV) population connects with the sympatric rearing grounds via relatively low salinity waters, whereas the San Pablo (SP) population must pass through the relatively high-salinity Upper SFE to reach its allopatric downstream spawning habitat. We hypothesize that if migration through SFE salinities to SP spawning grounds is more challenging for adult CV than SP splittail, then salinity tolerance, osmoregulatory capacity, and metabolic responses to salinity will differ between populations. Osmoregulatory disturbances, assessed by measuring plasma osmolality and ions, muscle moisture and Na + -K + -ATPase activity after 168 to 336 h at 11‰ salinity, showed evidence for a more robust osmoregulatory capacity in adult SP relative to CV splittail. While both resting and maximum metabolic rates were elevated in SP splittail in response to increased salinity, CV splittail metabolic rates were unaffected by salinity. Further, the calculated difference between resting and maximum metabolic values, aerobic scope, did not differ significantly between populations. Therefore, improved osmoregulation came at a metabolic cost for SP splittail but was not associated with negative impacts on scope for aerobic metabolism. These results suggest that SP splittail may be physiologically adjusted to allow for migration through higher-salinity waters. The trends in interpopulation variation in osmoregulatory and metabolic responses to salinity exposures support our hypothesis of greater salinity-related challenges to adult CV than SP splittail migration and are consistent with our previous findings for juvenile splittail populations, further supporting our recommendation of population-specific management.

California↗

Demographic estimation methods for plants with dormancy

Demographic studies in plants appear simple because unlike animals, plants do not run away. Plant individuals can be marked with, e.g., plastic tags, but often the coordinates of an individual may be sufficient to identify it. Vascular plants in temperate latitudes have a pronounced seasonal life–cycle, so most plant demographers survey their study plots once a year often during or shortly after flowering. Life–states are pervasive in plants, hence the results of a demographic study for an individual can be summarized in a familiar encounter history, such as 0VFVVF000. A zero means that an individual was not seen in a year and a letter denotes its state for years when it was seen aboveground. V and F here stand for vegetative and flowering states, respectively. Probabilities of survival and state transitions can then be obtained by mere counting. Problems arise when there is an unobservable dormant state, i.e., when plants may stay belowground for one or more growing seasons. Encounter histories such as 0VF00F000 may then occur where the meaning of zeroes becomes ambiguous. A zero can either mean a dead or a dormant plant. Various ad hoc methods in wide use among plant ecologists have made strong assumptions about when a zero should be equated to a dormant individual. These methods have never been compared among each other. In our talk and in Kéry et al. (submitted), we show that these ad hoc estimators provide spurious estimates of survival and should not be used. In contrast, if detection probabilities for aboveground plants are known or can be estimated, capturerecapture (CR) models can be used to estimate probabilities of survival and state–transitions and the fraction of the population that is dormant. We have used this approach in two studies of terrestrial orchids, Cleistes bifaria (Kéry et al., submitted) and Cypripedium reginae (Kéry & Gregg, submitted) in West Virginia, U.S.A. For Cleistes, our data comprised one population with a total of 620 marked ramets over 10 years, and for Cypripedium , two populations with 98 and 258 marked ramets over 11 years. We chose the ramet (= single stem or shoot) as the demographic unit of our study since there was no way distinguishing among genets (genet = genetical individual, i.e., the “individual” that animal ecologists are mostly concerned with). This will introduce some non–independence into the data, which can nevertheless be dealt with easily by correcting variances for overdispersion. Using ramets instead of genets has the further advantage that individuals can be assigned to a state such as flowering or vegetative in an unambiguous manner. This is not possible when genets are the demographic units. In all three populations, auxiliary data was available to show that detection probability of aboveground plants was m 0.995 We fitted multistate models in program MARK by specifying three states (D, V, F), even though the dormant state D does not occur in the encounter histories. Detection probability is fixed at 1 for the vegetative (V) and the flowering state (F) and at zero for the dormant state (D). Rates of survival and of state transitions as well as slopes of covariate relationships can be estimated and LRT or the AIC machinery be used to select among models. To estimate the fraction of the population in the unobservable dormant state, the encounter histories are collapsed to 0 (plant not observed aboveground) and 1 (plant observed aboveground). The Cormack–Jolly–Seber model without constraints on detection probability is used to estimate detection probability, the complement of which is the estimated fraction of the population in the dormant state. Parameter identifiability is an important issue in multi state models. We used the Catchpole–Morgan–Freeman approach to determine which parameters are estimable in principle in our multi state models. Most of 15 tested models were indeed estimable with the notable exception of the most general model, which has fully interactive state- and time-dependent survival and state transition rates. This model would become identifiable if at least some plants would be excavated in years when they do not show up aboveground. Our analyses for three analyzed populations of Cleistes and Cypripedium yielded annual ramet survival rates ranging from 0.86–0.96. Estimates of the average fraction dormant ranged from 0.02–0.30, but with up to half a population in the dormant state in some years. Ultrastructural modeling enables interesting hypotheses to be tested about the relationships of demographic rates with climatic covariates for instance. Such covariate modeling makes the CR approach particularly interesting for evolutionary–ecological questions about, e.g., the adaptive significance of the dormant state.

Animal Biodiversity and Conservation↗

Using environmental DNA (eDNA) to assess the presence of cavefish and cave crayfish populations in caves of the Ozark Highlands

Many cavefishes and cave crayfishes are considered of conservation concern; however, sampling these species is inherently difficult given their occupied environments. The goal of our project was to verify the presence of select karst organisms while developing the foundation for sampling approaches that might be useful to conservation and management agencies. Our project objectives were to develop assays to amplify deoxyribonucleic acid (DNA) from several species of Ozark cavefishes and cave crayfishes and complete an initial surveillance of locations across the Ozark Highlands using environmental DNA (eDNA). Using DNA either provided by agency cooperators or that we extracted from tissue samples, we PCR amplified and then sequenced the Cytochrome Oxidase 1 (CO1) gene for cave crayfishes and the NADH Dehydrogenase Subunit 2 (ND2) gene for cavefishes. We developed species-specific primers and probes for five cave crayfishes and two cavefishes. From February 2017 to May 2017, we sampled 1–5 sampling units from 42 caves, wells, and springs (i.e., sites) using eDNA and traditional visual surveys. We measured physicochemical parameters at each sampling unit to estimate detection probability associated with both techniques. We also calculated two occupancy covariates for each site using geospatial data. We successfully amplified Troglichthys rosae DNA from the environment and detected DNA representing this species at 24 of 40 sites. At 16 of the sites where we detected T. rosae DNA, we did not visually observe the species. Although our assay for Typlichthys eigenmanni successfully amplified the target DNA from the environment, it also resulted in false absences where the species was visually confirmed. Using eDNA to detect cave crayfishes was much more difficult. The assay for Cambarus subterraneus did not work for eDNA samples and we were unable to pick up DNA from the environment, even at locations where it was visually confirmed. Alternatively, the eDNA surveys worked well for C. tartarus and we were able to amplify DNA at every site where it was visually observed. Our assay for C. aculabrum was based on a single sample obtained from GenBank, and did not amplify eDNA from field samples. Lastly, our eDNA results from samples in the known range of Orconectes stygocaneyi suggested the species may be found at an additional cave. Detection using eDNA based on our O. stygocaneyi assay was likely low because it was designed from a pseudogene; however, positive eDNA samples were sequenced to confirm species-specific DNA. Detection probability of both cavefishes and cave crayfishes varied by survey technique and was influenced by water volume, water clarity, water velocity, and substrate. Detection of cavefishes and cave crayfishes via visual surveys decreased when water volume increased, whereas detection using eDNA increased with greater water volume. Detection between taxa using either sample method was highest in habitats classified by fine substrates, except for eDNA detection of crayfishes which was greatest in coarse substrates. Detection of cavefishes increased with water clarity, but detection of cave crayfishes increased with turbidity. Detection probability of both cavefishes and crayfishes using eDNA increased slightly with water velocity, but decreased with visual surveys as water velocity increased. Occupancy by both taxa was positively related to particular geologic series. Crayfish occupancy was negatively related to fine-scale anthropogenic disturbance (i.e., 500-m buffer around the site), whereas crayfish showed no relationship with disturbance. Our results suggest possible range extensions, provide insights to factors driving detection using both sample techniques, and suggest areas where recharge zones may be shared among caves. Future efforts focused on a comprehensive evaluation of genetic diversity among cave crayfishes to improve assay design could improve detection and the applicability of eDNA as a supplemental and non-invasive sampling approach.

Cooperator Science Series↗

Connectivity of Mojave Desert tortoise populations—Management implications for maintaining a viable recovery network

Executive Summary The historic distribution of Mojave desert tortoises ( Gopherus agassizii ) was relatively continuous across the range, and the importance of tortoise habitat outside of designated tortoise conservation areas (TCAs) to recovery has long been recognized for its contributions to supporting gene flow between TCAs and to minimizing impacts and edge effects within TCAs. However, connectivity of Mojave desert tortoise populations has become a concern because of recent and proposed development of large tracts of desert tortoise habitat that cross, fragment, and surround designated conservation areas. This paper summarizes the underlying concepts and importance of connectivity for Mojave desert tortoise populations by reviewing current information on connectivity and providing information to managers for maintaining or enhancing desert tortoise population connectivity as they consider future proposals for development and management actions. Maintaining an ecological network for the Mojave desert tortoise, with a system of core habitats (TCAs) connected by linkages, is necessary to support demographically viable populations and long-term gene flow within and between TCAs. There are four points for wildlife and land-management agencies to consider when making decisions that could affect connectivity of Mojave desert tortoise populations (for example, in updating actions in resource management plans or amendments that could help maintain or restore functional connectivity in light of the latest information): Management of all desert tortoise habitat for persistence and connectivity . Desert tortoise populations continue to decline within most TCAs, and it is unlikely that trends are better in populations outside protected areas. Fragmentation exacerbates negative population trends by breaking large continuous populations into smaller isolated populations. Connectivity within large populations can enhance resilience to localized disturbances due to rescue by neighboring individuals. In contrast, smaller fragmented populations are resistant to rescue by their isolation and thus could suffer irreversible declines to extirpation from a variety of threats and stochastic events. Enhanced threat reduction to reverse declines within TCAs and to maintain occupied habitat in the surrounding matrix would help reduce the variability in population growth rates and improve the resilience of protected populations even while implementing efforts to improve connectivity. Each TCA has unique strengths and weaknesses regarding its ability to support minimum sustainable populations based on areal extent and its ability to support population increases based on landscape connection with adjacent populations. Considering how proposed projects (inside or outside of TCAs) affect connectivity and the ability of TCAs to support at least 5,000 adult tortoises (the numerical goal for each TCA) could help managers to maintain the resilience of TCAs to population declines. The same project, in an alternative location, could have very different impacts on local and regional populations. For example, within the habitat matrix surrounding TCAs, narrowly delineated corridors may not allow for natural population dynamics if they do not accommodate overlapping home ranges along most of their widths so that tortoises reside, grow, find mates, and produce offspring that can replace older tortoises. In addition, most habitat outside TCAs may receive more surface disturbance than habitat within TCAs. Therefore, managing the entire remaining matrix of desert tortoise habitat for permeability may be better than delineating fixed corridors. These concepts apply, especially given uncertainty about long-term condition of habitat, within and outside of TCAs under a changing climate. Ultimately, questions such as “ What are the critical linkages that need to be protected ?” could be better framed as “ How can we manage the remaining habitat matrix in ways that sustain ecological processes and habitat suitability for special status species ?” Land-management decisions made in the context of the latter question may be more conducive to maintenance of a functional ecological network. Limitations on landscape-level disturbance across habitat managed for the desert tortoise Clearly delineating habitat linkages and differentiating them from non-delineated areas by the uses that are permitted or prohibited within them by specific management guidelines can help achieve functional connectivity. Such guidelines would be most effective if they considered and accounted for all surface disturbances (for example, temporary disturbances such as fiberoptic lines or off-highway vehicle routes, right-of-ways, utility-scale solar development, urbanization) to the extent possible. A weighted framework that varies with the permanence or severity of the disturbance, and can be additive to quantify cumulative effects, could be useful (Xiong, 2020). For example, minor roads can alter tortoise movements independently of other features (Peaden and others, 2017; Hromada and others, 2020), but if the isolated dirt road is accompanied by a powerline that encourages raven predation (Xiong, 2020), then the two features together may be additive. Ignoring minor or temporary disturbance on the landscape could result in a cumulatively large impact that is not explicitly acknowledged (Goble, 2009); therefore, understanding and quantifying all surface disturbance on a given landscape is prudent. In California, the Bureau of Land Management established 0.1–1.0 percent caps on new surface-disturbance for TCAs and mapped linkages that address the issues described in number 1 of this list. Nevada, Utah, and Arizona currently do not have surface-disturbance limits. Limits comparable to those in the Desert Renewable Energy Conservation Plan (DRECP) would be 0.5 percent within TCAs and 1 percent within the linkages modeled by Averill-Murray and others (2013). Limits in some areas of California within the Desert Renewable Energy Conservation Plan, such as Ivanpah Valley, are more restrictive, at 0.1 percent. Continuity across the state line in Nevada could be achieved with comparable limits in the adjacent portion of Ivanpah Valley, as well as the Greater Trout Canyon Translocation Area and the Stump Springs Regional Augmentation Site. These more restrictive limits would help protect remaining habitat in the major interstate connectivity pathway through Ivanpah Valley and focal areas of population augmentation that provide additional population connectivity along the western flank of the Spring Mountains. In a recent study that analyzed 13 years of desert tortoise monitoring data, nearly all desert tortoise observations were at sites in which 5 percent or less of the surrounding landscape within 1 kilometer was disturbed (Carter and others, 2020a). To help maintain tortoise habitability and permeability across all other non-conservation-designated tortoise habitat, all surface disturbance could be limited to less than 5-percent development per square kilometer because the 5-percent threshold for development is the point at which tortoise occupation drops precipitously (Carter and others, 2020a). However, although individual desert tortoises were observed at development levels up to 5 percent, we do not know the fitness or reproductive characteristics of these individuals. This level of development also may not allow for long-term persistence of healthy populations that are of adequate size needed for demographic or functional connectivity; therefore, a conservative interpretation suggests that, ideally, development could be lower. Lower development levels would be particularly useful in areas within the upper 5th percentile of connectivity values modeled by Gray and others (2019). Reducing ancillary threats in places where connectivity is restricted to narrow strips of habitat, for example, narrow mountain passes or vegetated strips between solar development, could enhance the functionality of these vulnerable linkages. In such areas, maintaining multiple, redundant linkages could further enhance overall connectivity. Minimization of mortality from roads and maximization of passage under roads . Roads pose a significant threat to the long-term persistence of local tortoise populations, and roads of high traffic volume lead to severe population declines, which ultimately fragments populations farther away from the roads. Three points (a.–c.) pertain to reducing direct mortality of tortoises on the many paved roads that cross desert tortoise habitat and to maintaining a minimal level of permeability across these roads: Tortoise-exclusion fencing tied into culverts, underpasses, overpasses, or other passages below roads in desert tortoise habitat, would limit vehicular mortality of tortoises and provide opportunities for movement across the roads. Installation of shade structures on the habitat side of fences installed in areas with narrow population-depletion zones would limit overheating of tortoises that may pace the fence. Passages below highways could be maintained or retrofitted to ensure safe tortoise access, for example, by filling eroded drop-offs or modifying erosion-control features such as rip-rap to make them safer and more passable for tortoises. Wildlife management agencies could work with transportation departments to develop construction standards that are consistent with hydrologic/erosion management goals, while also incorporating a design and materials consistent with tortoise survival and passage and make the standards widely available. The process would be most effective if the status of passages was regularly monitored and built into management plans. Healthy tortoise populations along fenced highways could be supported by ensuring that land inside tortoise-exclusion fences is not so degraded that it leads to degradation of tortoise habitat outside the exclusion areas. For example, severe invasive plant infestations inside a highway exclusion could cause an increase of invasive plants outside the exclusion area and degrade habitat; therefore, invasive plants inside road rights of way could be mown or treated with herbicide to limit their spread into adjacent tortoise habitat and minimize the risk of these plants carrying wildfires into adjacent habitat. Adaptation of management based on new information . Future research will continue to build upon and refine models related to desert tortoise population connectivity and develop new ones. New models could consider landscape levels of development and be constructed such that they share common foundations to support future synthesis efforts. If model development was undertaken in partnership with entities that are responsible for management of desert tortoise habitat, it would facilitate incorporation of current and future modeling results into their land management decisions. There are specific topics that may be clarified with further evaluation: The effects of climate change on desert tortoise habitat, distribution, and population connectivity; The effects of large-scale fires, especially within repeatedly burned habitat, on desert tortoise distribution and population connectivity; The ability of solar energy facilities or similar developments to support tortoise movement and presence by leaving washes intact; leaving native vegetation intact whenever possible, or if not possible, mowing the site, allowing vegetation to re-sprout, and managing weeds; and allowing tortoises to occupy the sites; and The design and frequency of underpasses necessary to maintain functional demographic and genetic connectivity across linear features, like highways.

Arizona, California, Nevada↗

Short-term impacts of a 4-lane highway on black bears in eastern North Carolina

Among numerous anthropogenic impacts on terrestrial landscapes, expanding transportation networks represent one of the primary challenges to wildlife conservation worldwide. Larger mammals may be particularly vulnerable because of typically low densities, low reproductive rates, and extensive movements. Although numerous studies have been conducted to document impacts of road networks on wildlife, inference has been limited because of experimental design limitations. During the last decade, the North Carolina Department of Transportation (NCDOT) rerouted and upgraded sections of United States Highway 64 between Raleigh and the Outer Banks to a 4-lane, divided highway. A new route was selected for a 24.1-km section in Washington County. The new section of highway included 3 wildlife underpasses with adjacent wildlife fencing to mitigate the effects of the highway on wildlife, particularly American black bears (Ursus americanus). We assessed the short-term impacts of the new highway on spatial ecology, population size, survival, occupancy, and gene flow of black bears. We tested our research hypotheses using a before-after control-impact (BACI) study design. We collected data during 2000–2001 (preconstruction phase) and 2006–2007 (postconstruction phase) in the highway project area and a nearby control area (each approx. 11,000 ha), resulting in 4 groups of data (i.e., pre- or postconstruction study phase, treatment or control area). We captured and radiocollared 57 bears and collected 5,775 hourly locations and 4,998 daily locations. Using mixed-model analysis of variance and logistic regression, we detected no differences in home ranges, movement characteristics, proximity to the highway alignment, or habitat use between the 2 study phases, although minimum detectable effect sizes were large for several tests. However, after completion of the new highway, bears on the treatment area became less inactive in morning, when highway traffic was low, compared with bears on the control area (F 1, 43 = 6.05, P = 0.018). We used DNA from hair samples to determine if population size and site occupancy decreased following highway construction. For each study phase, we collected black bear hair from 70 hair snares on each study area during 7 weekly sampling periods and generated genotypes using 10 microsatellite loci. We used the multilocus genotypes to obtain capture histories for 226 different bears and used capture-mark-recapture models to estimate population size. Model-averaged estimates of population size decreased on the treatment area from 87.7 bears before construction to 31.6 bears after construction (64% reduction) and on the control area from 163.6 bears to 108.2 bears (34% reduction). Permutation procedures indicated this reduction was proportionally greater for the treatment area (P = 0.086). We also applied a spatially explicit capture-recapture technique to test our research hypothesis. The model with the most support indicated a greater change in density on the treatment area (69% reduction) compared with the control area (24% reduction). We did not observe a treatment effect based on survival of radiocollared bears. We used bear visits to hair snares as detections in multi-season occupancy models and found that occupancy decreased more on the treatment area (preconstruction: Ψ = 0.84; postconstruction: Ψ = 0.44; 48% decline) than the control area (preconstruction: Ψ = 0.91; postconstruction: Ψ = 0.81; 11% decline), primarily as a function of a greater probability of site extinctions (ε) on the treatment area (ε = 0.57) than the control area (ε = 0.17). Finally, individual- and population-based analyses of contemporary gene flow did not indicate the highway was a barrier to movements. Black bear use of the 3 wildlife underpasses was infrequent (17 verified crossings based on remote cameras, track surveys, and telemetry). Only 4 of 8 bears with home ranges near the highway were documented crossing the highway (n = 36 crossings), of which 2 were killed in vehicle collisions. Six additional bears were killed in vehicle collisions from May 2007 to November 2008, after we completed field work. Harvest data indicated that hunting mortality alone could explain the population decline on the control area. On the treatment area, however, hunting mortality only accounted for an approximately 40% population decline; the additional 30% decline we observed likely was caused by other mortality. We speculate vehicle collisions were primarily responsible. We conclude that impacts of the new highway on resident black bears occurred at the population level, rather than the individual or genetic level, but that the impact was smaller than harvest mortality. Increased activity by remaining bears when traffic volumes were low indicated behavioral plasticity. Bear use of the underpasses seemed sufficient to maintain gene flow between areas north and south of the new highway. Effectiveness of wildlife underpasses to reduce mortality of black bears may be enhanced if mitigation includes continuous fencing between crossing structures. For small, isolated populations of threatened or endangered large mammals, the potential demographic impacts of highways are an essential consideration in the transportation planning process. Control of mortality factors and maintaining demographic connectivity are particularly important.

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↗