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A geostatistical state‐space model of animal densities for stream networks

Population dynamics are often correlated in space and time due to correlations in environmental drivers as well as synchrony induced by individual dispersal. Many statistical analyses of populations ignore potential autocorrelations and assume that survey methods (distance and time between samples) eliminate these correlations, allowing samples to be treated independently. If these assumptions are incorrect, results and therefore inference may be biased and uncertainty underestimated. We developed a novel statistical method to account for spatiotemporal correlations within dendritic stream networks, while accounting for imperfect detection in the surveys. Through simulations, we found this model decreased predictive error relative to standard statistical methods when data were spatially correlated based on stream distance and performed similarly when data were not correlated. We found that increasing the number of years surveyed substantially improved the model accuracy when estimating spatial and temporal correlation coefficients, especially from 10 to 15 yr. Increasing the number of survey sites within the network improved the performance of the nonspatial model but only marginally improved the density estimates in the spatiotemporal model. We applied this model to brook trout data from the West Susquehanna Watershed in Pennsylvania collected over 34 yr from 1981 to 2014. We found the model including temporal and spatiotemporal autocorrelation best described young of the year (YOY) and adult density patterns. YOY densities were positively related to forest cover and negatively related to spring temperatures with low temporal autocorrelation and moderately high spatiotemporal correlation. Adult densities were less strongly affected by climatic conditions and less temporally variable than YOY but with similar spatiotemporal correlation and higher temporal autocorrelation.

Ecological Applications↗

Differences in ecosystem carbon distribution and nutrient cycling linked to forest tree species composition in a mid-successional boreal forest

In the boreal forest of Alaska, increased fire severity associated with climate change is expanding deciduous forest cover in areas previously dominated by black spruce ( Picea mariana ). Needle-leaf conifer and broad-leaf deciduous species are commonly associated with differences in tree growth, carbon (C) and nutrient cycling, and C accumulation in soils. Although this suggests that changes in tree species composition in Alaska could impact C and nutrient pools and fluxes, few studies have measured these linkages. We quantified C, nitrogen, phosphorus, and base cation pools and fluxes in three stands of black spruce and Alaska paper birch ( Betula neoalaskana ) that established following a single fire event in 1958. Paper birch consistently displayed characteristics of more rapid C and nutrient cycling, including greater aboveground net primary productivity, higher live foliage and litter nutrient concentrations, and larger ammonium and nitrate pools in the soil organic layer (SOL). Ecosystem C stocks (aboveground + SOL + 0–10 cm mineral soil) were similar for the two species; however, in black spruce, 78% of measured C was found in soil pools, primarily in the SOL, whereas aboveground biomass dominated ecosystem C pools in birch forest. Radiocarbon analysis indicated that approximately one-quarter of the black spruce SOL C accumulated prior to the 1958 fire, whereas no pre-fire C was observed in birch soils. Our findings suggest that tree species exert a strong influence over C and nutrient cycling in boreal forest and forest compositional shifts may have long-term implications for ecosystem C and nutrient dynamics.

Ecosystems↗

Seasonal precipitation and soil moisture relationships across forests and woodlands in the southwestern United States

Precipitation [P: mm] controls forest and woodland dynamics in the southwestern United States (SWUS) by altering soil moisture [ θ : mm 3 mm −3 ] availability, but the influence of P on θ is complex, varying across space and time. We evaluated seasonal P and θ relationships at shallow (0‐20 cm) and intermediate (50 cm) soil depths for 9 semiarid forest and woodland sites (56 total years), which comprised 3 elevation gradients in the SWUS. We developed time‐varying definitions of winter (snow accumulation), spring (moisture recharge), and summer (moisture deficit), and determined how these sites exhibited similar P influence on θ across depths in the soil profile, between seasons, and in seasons with above‐ and below‐average P. Higher elevation sites ( > 2800 m) experienced greater winter P, longer springs, and shorter summers compared to lower elevation sites ( < 2500 m). Seasons with above‐ and below‐average P reduced elevation‐associated differences. θ at 0‐20 cm was generally decoupled from θ at 50 cm in seasons with average and below‐average P, imparted by differences in spring and summer rainfall versus winter snowfall. Notably, across‐season influence of θ (e.g. a season's similarity to subsequent seasons) was high when the first season experienced above‐ or below‐average P, and the subsequent season experienced average P, illustrating an important temporal connection initiated by wet and dry conditions. These results illustrate similarities in P‐ θ relationships across widely differing ecosystems in the SWUS, and elucidate how these relationships may be altered in a changing climate.

Journal of Geophysical Research Biogeosciences↗

Improved arrival-date estimates of Arctic-breeding Dunlin ( Calidris alpina arcticola )

The use of stable isotopes in animal ecology depends on accurate descriptions of isotope dynamics within individuals. The prevailing assumption that laboratory-derived isotopic parameters apply to free-living animals is largely untested. We used stable carbon isotopes (&delta; 13 C) in whole blood from migratory Dunlin ( Calidris alpina arcticola ) to estimate an in situ turnover rate and individual diet-switch dates. Our in situ results indicated that turnover rates were higher in free-living birds, in comparison to the results of an experimental study on captive Dunlin and estimates derived from a theoretical allometric model. Diet-switch dates from all 3 methods were then used to estimate arrival dates to the Arctic; arrival dates calculated with the in situ turnover rate were later than those with the other turnover-rate estimates, substantially so in some cases. These later arrival dates matched dates when local snow conditions would have allowed Dunlin to settle, and agreed with anticipated arrival dates of Dunlin tracked with light-level geolocators. Our study presents a novel method for accurately estimating arrival dates for individuals of migratory species in which return dates are difficult to document. This may be particularly appropriate for species in which extrinsic tracking devices cannot easily be employed because of cost, body size, or behavioral constraints, and in habitats that do not allow individuals to be detected easily upon first arrival. Thus, this isotopic method offers an exciting alternative approach to better understand how species may be altering their arrival dates in response to changing climatic conditions.

The Auk↗

Lateral and subsurface flows impact arctic coastal plain lake water budgets

Arctic thaw lakes are an important source of water for aquatic ecosystems, wildlife, and humans. Many recent studies have observed changes in Arctic surface waters related to climate warming and permafrost thaw; however, explaining the trends and predicting future responses to warming is difficult without a stronger fundamental understanding of Arctic lake water budgets. By measuring and simulating surface and subsurface hydrologic fluxes, this work quantified the water budgets of three lakes with varying levels of seasonal drainage, and tested the hypothesis that lateral and subsurface flows are a major component of the post-snowmelt water budgets. A water budget focused only on post-snowmelt surface water fluxes (stream discharge, precipitation, and evaporation) could not close the budget for two of three lakes, even when uncertainty in input parameters was rigorously considered using a Monte Carlo approach. The water budgets indicated large, positive residuals, consistent with up to 70% of mid-summer inflows entering lakes from lateral fluxes. Lateral inflows and outflows were simulated based on three processes; supra-permafrost subsurface inflows from basin-edge polygonal ground, and exchange between seasonally drained lakes and their drained margins through runoff and evapotranspiration. Measurements and simulations indicate that rapid subsurface flow through highly conductive flowpaths in the polygonal ground can explain the majority of the inflow. Drained lakes were hydrologically connected to marshy areas on the lake margins, receiving water from runoff following precipitation and losing up to 38% of lake efflux to drained margin evapotranspiration. Lateral fluxes can be a major part of Arctic thaw lake water budgets and a major control on summertime lake water levels. Incorporating these dynamics into models will improve our ability to predict lake volume changes, solute fluxes, and habitat availability in the changing Arctic.

Alaska↗

Evaluating fish rescue as a drought adaptation strategy using a life cycle modeling approach for imperiled coho salmon

Projected intensification of drought as a result of climate change may reduce the capacity of streams to rear fish, exacerbating the challenge of recovering salmonid populations listed under the Endangered Species Act. Without management intervention, some stocks will likely go extinct as stream drying and fragmentation reduce juvenile survival to unsustainable levels. To offset drought‐related mortality, fish rescue programs have proliferated, whereby juvenile salmonids are captured and transferred to off‐site rearing facilities. However, the efficacy of this potential conservation tool remains poorly understood. We developed a life cycle model to examine the implications of fish rescue on the abundance of Coho Salmon Oncorhynchus kisutch across serial life stages. The simulation model examines scenarios with varying quantities of rescued fish, time in captivity, drought severity, and reduced smolt‐to‐adult return rates. Our results indicate that fish rescue can increase the abundance of adults and lower extinction risk, particularly for fish held in captivity for a full year. However, fish rescue can also decrease the abundance of adults and increase extinction risk if fish are held only for summer and there is limited winter habitat. We found that when fish rescue did increase returns, it functioned more like a stock enhancement program than a drought mitigation tool and it would likely lead to consecutive generations of captive rearing, which has been shown to have negative effects on fitness. We translated our model into an R Shiny application ( https://shiny.wdfw‐fish.us/CohoPopulationDynamics/ ) that allows users to explore how fish rescue affects Coho Salmon population dynamics through customized parameterization of the model to represent different systems or different assumptions about the effects of fish rescue.

North American Journal of Fisheries Management↗

Meeting ecological and societal needs for freshwater

Human society has used freshwater from rivers, lakes, groundwater, and wetlands for many different urban, agricultural, and industrial activities, but in doing so has overlooked its value in supporting ecosystems. Freshwater is vital to human life and societal well-being, and thus its utilization for consumption, irrigation, and transport has long taken precedence over other commodities and services provided by freshwater ecosystems. However, there is growing recognition that functionally intact and biologically complex aquatic ecosystems provide many economically valuable services and long-term benefits to society. The short-term benefits include ecosystem goods and services, such as food supply, flood control, purification of human and industrial wastes, and habitat for plant and animal life—and these are costly, if not impossible, to replace. Long-term benefits include the sustained provision of those goods and services, as well as the adaptive capacity of aquatic ecosystems to respond to future environmental alterations, such as climate change. Thus, maintenance of the processes and properties that support freshwater ecosystem integrity should be included in debates over sustainable water resource allocation. The purpose of this report is to explain how the integrity of freshwater ecosystems depends upon adequate quantity, quality, timing, and temporal variability of water flow. Defining these requirements in a comprehensive but general manner provides a better foundation for their inclusion in current and future debates about allocation of water resources. In this way the needs of freshwater ecosystems can be legitimately recognized and addressed. We also recommend ways in which freshwater ecosystems can be protected, maintained, and restored. Freshwater ecosystem structure and function are tightly linked to the watershed or catchment of which they are a part. Because riverine networks, lakes, wetlands, and their connecting groundwaters, are literally the “sinks” into which landscapes drain, they are greatly influenced by terrestrial processes, including many human uses or modifications of land and water. Freshwater ecosystems, whether lakes, wetlands, or rivers, have specific requirements in terms of quantity, quality, and seasonality of their water supplies. Sustainability normally requires these systems to fluctuate within a natural range of variation. Flow regime, sediment and organic matter inputs, thermal and light characteristics, chemical and nutrient characteristics, and biotic assemblages are fundamental defining attributes of freshwater ecosystems. These attributes impart relatively unique characteristics of productivity and biodiversity to each ecosystem. The natural range of variation in each of these attributes is critical to maintaining the integrity and dynamic potential of aquatic ecosystems; therefore, management should allow for dynamic change. Piecemeal approaches cannot solve the problems confronting freshwater ecosystems. Scientific definitions of the requirements to protect and maintain aquatic ecosystems are necessary but insufficient for establishing the appropriate distribution between societal and ecosystem water needs. For scientific knowledge to be implemented science must be connected to a political agenda for sustainable development. We offer these recommendations as a beginning to redress how water is viewed and managed in the United States: (1) Frame national and regional water management policies to explicitly incorporate freshwater ecosystem needs, particularly those related to naturally variable flow regimes and to the linking of water quality with water quantity; (2) Define water resources to include watersheds, so that freshwaters are viewed within a landscape, or systems context; (3) Increase communication and education across disciplines, especially among engineers, hydrologists, economists, and ecologists to facilitate an integrated view of freshwater resources; (4) Increase restoration efforts, using well-grounded ecological principles as guidelines; (5) Maintain and protect the remaining freshwater ecosystems that have high integrity; and (6) Recognize the dependence of human society on naturally functioning ecosystems.

Ecological Applications↗

Water-quality conditions in Upper Klamath Lake, Oregon, 2002-04

Eleven (2002) to 14 (2003 and 2004) continuous water-quality monitors that measured pH, dissolved oxygen, temperature, and specific conductance, were placed in Upper Klamath Lake to support a telemetry tracking study of endangered adult shortnose and Lost River suckers. Samples for the analysis of chlorophyll a and nutrients were collected at a subset of the water-quality monitor sites in each year. The seasonal pattern in the occurrence of supersaturated dissolved oxygen concentrations and high pH associated with photosynthetic activity, as well as the undersaturated dissolved oxygen concentrations associated with oxygen demand through respiration and decay in excess of photosynthetic production, were well described by the dynamics of the massive blooms of Aphanizomenon flos aquae (AFA) that occur each year. Data from the continuous monitors provided a means to quantify the occurrence, duration, and spatial extent of water-quality conditions potentially harmful to fish (dissolved- oxygen concentration less than 4 milligrams per liter, pH greater than 9.7, and temperature greater than 28 degrees Celsius) in the northern part of the lake, where the preferred adult sucker habitat is found. There were few observations of temperature greater than 28 degrees Celsius, suggesting that temperature is not a significant source of chronic stress to fish, although its role in the spread of disease is harder to define. Observations of pH greater than 9.7 were common during times when the AFA bloom was growing rapidly, so pH may be a source of chronic stress to fish. Dissolved oxygen concentrations less than 4 milligrams per liter were common in all 3 years at the deeper sites, in the lower part of the water column and for short periods during the day. Less common were instances of widespread low dissolved oxygen, throughout the water column and persisting through the entire day, but this was the character of a severe low dissolved oxygen event (LDOE) that culminated in the start of a fish die-off in 2003. Documented evidence indicates that LDOEs played a role in three fish die-offs in the mid-1990s as well. In the historical context of 15 years of climate and water-quality data, 3 out of 4 of the recent fish die-off years, 1996, 1997, and 2003, were characterized by low winds and high temperatures in July or August coincident with the start of the die-off. High temperatures accelerate the oxygen demanding processes that lead to a LDOE. The role of low winds remains inconclusive, but it could include the development of stratification in the water column and/or the alteration of the wind-driven circulation pattern. At a site centrally located in the study area, die-off years could be successfully identified in the historical data by screening for water characterized by exceptionally low chlorophyll a concentration, exceptionally low dissolved oxygen concentration throughout the water column (not just near the bottom), and exceptionally high ammonia concentration and water temperature, just prior to or coincident with the start of a fish die-off. These conditions indicate that a severe decline in the AFA bloom and conversion of most of the organic matter into inorganic form had taken place.

Scientific Investigations Report↗

Evidence of counter-gradient growth in western pond turtles ( Actinemys marmorata ) across thermal gradients

Counter-gradient growth, where growth per unit temperature increases as temperature decreases, can reduce the variation in ectothermic growth rates across environmental gradients. Understanding how ectothermic species respond to changing temperatures is essential to their conservation and management due to human-altered habitats and changing climates. Here, we use two contrasting populations of western pond turtles ( Actinemys marmorata ) to model the effect of artificial and variable temperature regimes on growth and age at reproductive maturity. The two populations occur on forks of the Trinity River in northern California, U.S.A. The South Fork Trinity River (South Fork) is unregulated, while the main stem of the Trinity River (Main Stem) is dammed and has peak seasonal temperatures that are approximately 10 &deg;C colder than the South Fork. Consistent with other studies, we found reduced annual growth rates for turtles in the colder Main Stem compared to the warmer South Fork. The South Fork population matured approximately 9 year earlier, on average, and at a larger body size than the Main Stem population. When we normalised growth rates for the thermal opportunity for growth using water-growing degree-days (GDD), we found the reverse for growth rates and age at reproductive maturity. Main Stem turtles grew approximately twice as fast as South Fork turtles per GDD. Main Stem turtles also required approximately 50% fewer GDD to reach their smaller size at reproductive maturity compared to the larger South Fork turtles. We found we could accurately hindcast growth rates based on water temperatures estimated from the total volume of discharge from the dam into the Main Stem, providing a management tool for predicting the impacts of the dam on turtle growth rates. Given the importance of size and age at reproductive maturity to population dynamics, this information on counter-gradient growth will improve our ability to understand and predict the consequences of dam operations for downstream turtle populations.

California↗

Forestry practices and aquatic biodiversity: Fish

In the Pacific Northwest, fish communities are found in a diverse array of aquatic habitats ranging from the large coastal rivers of the temperate rainforests, to the fragmented and sometimes ephemeral streams of the xeric interior basins, and high-elevation streams and lakes in the mountainous areas (Rieman et al. 2003). Only high-elevation lakes and streams isolated above barriers to fish passage remained historically devoid of fish because they were never invaded following Pleistocene glaciation (Smith 1981). Despite this widespread distribution and once great population abundances, taxonomic diversity of fishes in these forested systems is naturally lower than in aquatic habitats in the eastern U.S. (Reeves, Bisson, and Dambacher 1998). Interactions among factors that influence species richness in aquatic systems (e.g., basin size, long-term stability of habitat, and barriers to colonization; Smith 1981) continue to influence the occurrence and persistence of fishes in these systems today. Consequently, the larger low-elevation rivers and estuaries support the greatest variety of fish species. In the high-elevation tributary streams, fish communities are less complex because these aquatic systems were less climatically and geologically stable, and fish populations were smaller and more prone to local extirpation. Furthermore, barriers to fish passage inhibited dispersal and colonization (Smith 1981). Streams in forested landscapes generally support salmon and trout, Oncorhynchus spp., whitefish Prosopium spp., sculpins Cottus spp., suckers Catostomus spp., and minnows (Cyprinidae), but in some of the colder streams, chars (e.g., Salvelinus confluentus and Salvelinus malma ) and lampreys (Petromyzontidae)may also occur (Rieman et al. 2003). Although biodiversity defined in terms of fish species richness is low in the Pacific Northwest, intraspecific variability is high, and polytypic fish species are common in the diverse aquatic habitats of the region. For example, the salmonids in the coastal rivers and streams, and the larger interconnected streams, rivers, and lakes of the interior exhibit a variety of ecotypes and migratory life histories (Healey 1986; Trotter 1989; Larson and McIntire 1993; Northcote 1997). This life-history variation appears to be associated with adaptation to spatial and temporal variation in environment (e.g., Schaffer and Elson 1975; Carl and Healey 1984; Beacham and Murray 1987), and there is some evidence of the genetic heritability of life-history traits (Carl and Healey 1984; Gharrett and Smoker 1993; Hankin, Nicholas, and Downey 1993). Persistence of any level of biological organization (e.g., life-history type, population, metapopulation, subspecies, species, community) is related to the interaction of environmental and biological components, and intraspecific diversity is a means of spreading risk ( sensu den Boer 1968) of extirpation in dynamic environments (Gresswell 1999). Unfortunately, despite the broad distribution and extensive intraspecific diversity, persistence of native fishes is uncertain in the Pacific Northwest. Many populations of anadromous salmonids, once synonymous with vigorous biological communities throughout the region, are threatened with extinction (Nehlsen, Williams, and Lichatowich 1991; Frissell 1993; Thurow, Lee, and Rieman 1997). Furthermore, over half of the native taxa in the Columbia River Basin are either listed under the Endangered Species Act, are being considered for listing, or are deemed sensitive by the management agencies (Lee et al. 1997; Thurow, Lee, and Rieman 1997). Potamodromous species like bull trout Salvelinus confluentus are estimated to occur as strong populations in less than 5% of their potential range (Rieman, Lee, and Thurow 1997). Although not currently listed under the endangered species list, the coastal cutthroat trout Oncorhynchus clarki is managed as a sensitive species in Oregon and California (Hall, Bisson, and Gresswell 1997). Native non-game fishes have rarely been monitored, but populations of species such as large-scale suckers ( Catostomus macrocheilus ), squawfish ( Ptychocheilus umpquae ), and Pacific lamprey ( Lampetra tridentata ) also are declining in some drainages (Oregon Department of Fish and Wildlife, unpublished data).

Pacific Northwest↗

Effect of tree line advance on carbon storage in NW Alaska

We investigated the size, distribution, and temporal dynamics of ecosystem carbon (C) pools in an area of recent tree line advance, northwest Alaska. Repeat aerial photographs show forest cover increased ???10% in our study area since 1949. We sampled C pools of four principal ecosystem types, tussock tundra, shrub tundra, woodland, and forest, all located on a 600-800 year old river terrace. Significant differences between ecosystem C pools, both above ground and below ground existed. Tundra sites store >22.2 kg C/m2, shrub tundra sites and woodland sites store 9.7 kg C/m2 and 14.3 kg C/m2, respectively, and forest sites store 14.4 kg C/m2. Landscape variation of total ecosystem C was primarily due to organic soil C and was secondarily due to C stored in trees. Soil C/N profiles of shrub tundra sites and woodland sites showed similarities with forest site soils at surface and tundra site soils at depth. We hypothesize that tundra systems transformed to forest systems in this area under a progression of permafrost degradation and enhanced drainage. On the basis of C pool estimates for the different ecosystem types, conversion of tundra sites to forest may have resulted in a net loss of > 7.8 kg C/m2, since aboveground C gains were more than offset by belowground C losses to decomposition in the tundra sites. Tree line advance therefore might not increase C storage in high-latitude ecosystems and thus might not, as previously suggested, act as a negative feedback to warming. Key to this hypothesis and to its projection to future climate response is the fate of soil carbon upon warming and permafrost drainage. Copyright 2006 by the American Geophysical Union.

Journal of Geophysical Research G: Biogeosciences↗

Breaking down Palila decline: Assessing the role of drought and vegetation health in the population loss of an endangered Hawaiian honeycreeper

The Palila ( Loxioides bailleui ), the last member of the once speciose finch-billed Hawaiian honeycreeper clade (Drepanidinae) in the main Hawaiian Islands, faces critical conservation challenges as an endangered species. Understanding the drivers of its decline is essential for effective management. We used additive decomposition models to examine temporal trends in climatic variables (temperature, precipitation, drought) and Normalized Difference Vegetation Index (NDVI), a vegetation health metric hypothesized to be associated with long-term trends in Palila abundance at landscape (250 m) scales on the Island of Hawai'i. A breakpoint analysis identified 2005–2009 as critical years of Palila decline. Vegetation health metrics at the 250 m scale lined up well both spatially and temporally with trends in Palila declines, with a significant browning from January 2004 to January 2014. Given the strong correlation between vegetation health and drought metrics at the landscape scale (r = 0.75, p < 0.001), NDVI changes appeared driven by drought. To enable the future projection of habitat quality in this area, we explored a stepwise linear regression to explain the variation in MODIS NDVI in recent years. We found that 87 % of the variability in NDVI can be explained by wet season precipitation and vapor pressure deficit from the previous dry season. The model is largely driven by a strong positive correlation between wet season precipitation and NDVI (r = 0.72, adjusted p < 0.001). Areas that maintained a low likelihood of NDVI decline throughout the time series and experienced increases in predicted Palila count represent potential drought microrefugia for the species. This higher elevation microrefugia is likely resilient against decreases in wet season precipitation through supplemental water retention from fog drip. While NDVI rebounded after 2014, Palila have not recovered. Our analysis highlights the importance of trend decomposition for monitoring endangered species with limited rebound potential due to small population dynamics and indicate continued warm, dry conditions may prevent Palila recovery without intervention.

Hawaii↗

Effects of El Niño on distribution and reproductive performance of Black Brant

Climate in low-latitude wintering areas may influence temperate and high-latitude breeding populations of birds, but demonstrations of such relationships have been rare because of difficulties in linking wintering with breeding populations. We used long-term aerial surveys in Mexican wintering areas and breeding areas in Alaska, USA, to assess numbers of Black Brant ( Branta bernicla nigricans ; hereafter brant) on their principal wintering and breeding area in El Niño and non-El Niño years. We used Pollock's robust design to directly estimate probability of breeding and apparent annual survival of individually marked brant at the Tutakoke River (TR) colony, Alaska, in each year between 1988 and 2001. Fewer brant wintered in Mexico during every El Niño event since 1965. Fewer brant were observed on the principal breeding area following each El Niño since surveys began in 1985. Probability of breeding was negatively related to January sea surface temperature along the subtropical coast of North America during the preceding winter. Between 23% (five-year-olds or older) and 30% (three-year-olds) fewer brant nested in 1998 following the strong El Niño event in the winter of 1997–1998 than in non-El Niño years. This finding is consistent with life history theory, which predicts that longer-lived species preserve adult survival at the expense of reproduction. Oceanographic conditions off Baja California, apparently by their effect on Zostera marina (eelgrass), strongly influence winter distribution of brant geese and their reproduction (but not survival), which in turn affects ecosystem dynamics in Alaska.

Alaska↗

A modeling framework for integrated harvest and habitat management of North American waterfowl: Case-study of northern pintail metapopulation dynamics

We developed and evaluated the performance of a metapopulation model enabling managers to examine, for the first time, the consequences of alternative management strategies involving habitat conditions and hunting on both harvest opportunity and carrying capacity (i.e., equilibrium population size in the absence of harvest) for migratory waterfowl at a continental scale. Our focus is on the northern pintail ( Anas acuta ; hereafter, pintail), which serves as a useful model species to examine the potential for integrating waterfowl harvest and habitat management in North America. We developed submodel structure capturing important processes for pintail populations during breeding, fall migration, winter, and spring migration while encompassing spatial structure representing three core breeding areas and two core nonbreeding areas. A number of continental-scale predictions from our baseline parameterization (e.g., carrying capacity of 5.5 million, equilibrium population size of 2.9 million and harvest rate of 12% at maximum sustained yield [MSY]) were within 10% of those from the pintail harvest strategy under current use by the U.S. Fish and Wildlife Service. To begin investigating the interaction of harvest and habitat management, we examined equilibrium population conditions for pintail at the continental scale across a range of harvest rates while perturbing model parameters to represent: (1) a 10% increase in breeding habitat quality in the Prairie Pothole population (PR); and (2) a 10% increase in nonbreeding habitat quantity along in the Gulf Coast (GC). Based on our model and analysis, a greater increase in carrying capacity and sustainable harvest was seen when increasing a proxy for habitat quality in the Prairie Pothole population. This finding and underlying assumptions must be critically evaluated, however, before specific management recommendations can be made. To make such recommendations, we require (1) extended, refined submodels with additional parameters linking influences of habitat management and environmental conditions to key life-history parameters; (2) a formal sensitivity analysis of the revised model; (3) an integrated population model that incorporates empirical data for estimating key vital rates; and (4) cost estimates for changing these additional parameters through habitat management efforts. We foresee great utility in using an integrated modeling approach to predict habitat and harvest management influences on continental-scale population responses while explicitly considering putative effects of climate change. Such a model could be readily adapted for management of many habitat-limited species.

Ecological Modelling↗

Salt marsh ecosystem restructuring enhances elevation resilience and carbon storage during accelerating relative sea-level rise

Salt marshes respond to sea-level rise through a series of complex and dynamic bio-physical feedbacks. In this study, we found that sea-level rise triggered salt marsh habitat restructuring, with the associated vegetation changes enhancing salt marsh elevation resilience. A continuous record of marsh elevation relative to sea level that includes reconstruction of high-resolution, sub-decadal, marsh elevation over the past century, coupled with a lower-resolution 1500-year record, revealed that relative sea-level rose 1.5 ± 0.4 m, following local glacial isostatic adjustment (1.2 mm/yr). As sea-level rise has rapidly accelerated, the high marsh zone dropped 11 cm within the tidal frame since 1932, leading to greater inundation and a shift to flood- and salt-tolerant low marsh species. Once the marsh platform fell to the elevation favored by low-marsh Spartina alterniflora , the elevation stabilized relative to sea level. Currently low marsh accretion keeps pace with sea-level rise, while present day high marsh zones that have not transitioned to low marsh have a vertical accretion deficit. Greater biomass productivity, and an expanding subsurface accommodation space favorable for salt marsh organic matter preservation, provide a positive feed-back between sea-level rise and marsh platform elevation. Carbon storage was 46 ± 28 g C/m 2 /yr from 550 to 1800 CE, increasing to 129 ± 50 g C/m 2 /yr in the last decade. Enhanced carbon storage is controlled by vertical accretion rates, rather than soil carbon density, and is a direct response to anthropogenic eustatic sea-level rise, ultimately providing a negative feedback on climate warming.

Massachusetts↗

Quantifying drivers of wild pig movement across multiple spatial and temporal scales

Background The movement behavior of an animal is determined by extrinsic and intrinsic factors that operate at multiple spatio-temporal scales, yet much of our knowledge of animal movement comes from studies that examine only one or two scales concurrently. Understanding the drivers of animal movement across multiple scales is crucial for understanding the fundamentals of movement ecology, predicting changes in distribution, describing disease dynamics, and identifying efficient methods of wildlife conservation and management. Methods We obtained over 400,000 GPS locations of wild pigs from 13 different studies spanning six states in southern U.S.A., and quantified movement rates and home range size within a single analytical framework. We used a generalized additive mixed model framework to quantify the effects of five broad predictor categories on movement: individual-level attributes, geographic factors, landscape attributes, meteorological conditions, and temporal variables. We examined effects of predictors across three temporal scales: daily, monthly, and using all data during the study period. We considered both local environmental factors such as daily weather data and distance to various resources on the landscape, as well as factors acting at a broader spatial scale such as ecoregion and season. Results We found meteorological variables (temperature and pressure), landscape features (distance to water sources), a broad-scale geographic factor (ecoregion), and individual-level characteristics (sex-age class), drove wild pig movement across all scales, but both the magnitude and shape of covariate relationships to movement differed across temporal scales. Conclusions The analytical framework we present can be used to assess movement patterns arising from multiple data sources for a range of species while accounting for spatio-temporal correlations. Our analyses show the magnitude by which reaction norms can change based on the temporal scale of response data, illustrating the importance of appropriately defining temporal scales of both the movement response and covariates depending on the intended implications of research (e.g., predicting effects of movement due to climate change versus planning local-scale management). We argue that consideration of multiple spatial scales within the same framework (rather than comparing across separate studies post-hoc) gives a more accurate quantification of cross-scale spatial effects by appropriately accounting for error correlation.

Movement Ecology↗

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↗

Distribution of a climate-sensitive species at an interior range margin

Advances in understanding the factors that limit a species’ range, particularly in the context of climate change, have come disproportionately through investigations at range edges or margins. The margins of a species’ range might often correspond with anomalous microclimates that confer habitat suitability where the species would otherwise fail to persist. We addressed this hypothesis using data from an interior, climatic range margin of the American pika ( Ochotona princeps ), an indicator of relatively cool, mesic climates in rocky habitats of western North America. Pikas in Lava Beds National Monument, northeastern California, USA, occur at elevations much lower than predicted by latitude and longitude. We hypothesized that pika occurrence within Lava Beds would be associated primarily with features such as “ice caves” in which sub-surface ice persists outside the winter months. We used data loggers to monitor sub-surface temperatures at cave entrances and at non-cave sites, confirming that temperatures were cooler and more stable at cave entrances. We surveyed habitat characteristics and evidence of pika occupancy across a random sample of cave and non-cave sites over a 2-yr period. Pika detection probability was high (~0.97), and the combined occupancy of cave and non-cave sites varied across the 2 yr from 27% to 69%. Contrary to our hypothesis, occupancy was not higher at cave sites. Vegetation metrics were the best predictors of site use by pikas, followed by an edge effect and elevation. The importance of vegetation as a predictor of pika distribution at this interior range margin is congruent with recent studies from other portions of the species’ range. However, we caution that vegetation composition depends on microclimate, which might be the proximal driver of pika distribution. The microclimates available in non-cave crevices accessible to small animals have not been characterized adequately for lava landscapes. We advocate innovation in the acquisition and use of microclimatic data for understanding the distributions of many taxa. Appropriately scaled microclimatic data are increasingly available but rarely used in studies of range dynamics.

Ecosphere↗