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Leveraging constraints and biotelemetry data to pinpoint repetitively used spatial features

Satellite telemetry devices collect valuable information concerning the sites visited by animals, including the location of central places like dens, nests, rookeries, or haul‐outs. Existing methods for estimating the location of central places from telemetry data require user‐specified thresholds and ignore common nuances like measurement error. We present a fully model‐based approach for locating central places from telemetry data that accounts for multiple sources of uncertainty and uses all of the available locational data. Our general framework consists of an observation model to account for large telemetry measurement error and animal movement, and a highly flexible mixture model specified using a Dirichlet process to identify the location of central places. We also quantify temporal patterns in central place use by incorporating ancillary behavioral data into the model; however, our framework is also suitable when no such behavioral data exist. We apply the model to a simulated data set as proof of concept. We then illustrate our framework by analyzing an Argos satellite telemetry data set on harbor seals ( Phoca vitulina ) in the Gulf of Alaska, a species that exhibits fidelity to terrestrial haul‐out sites.

Alaska↗

Monitoring dynamic spatio-temporal ecological processes optimally

Population dynamics vary in space and time. Survey designs that ignore these dynamics may be inefficient and fail to capture essential spatio‐temporal variability of a process. Alternatively, dynamic survey designs explicitly incorporate knowledge of ecological processes, the associated uncertainty in those processes, and can be optimized with respect to monitoring objectives. We describe a cohesive framework for monitoring a spreading population that explicitly links animal movement models with survey design and monitoring objectives. We apply the framework to develop an optimal survey design for sea otters in Glacier Bay. Sea otters were first detected in Glacier Bay in 1988 and have since increased in both abundance and distribution; abundance estimates increased from 5 otters to >5,000 otters, and they have spread faster than 2.7 km/yr. By explicitly linking animal movement models and survey design, we are able to reduce uncertainty associated with forecasting occupancy, abundance, and distribution compared to other potential random designs. The framework we describe is general, and we outline steps to applying it to novel systems and taxa.

Ecology↗

Ecological consequences of anomalies in atmospheric moisture and snowpack

Although increased frequency of extreme‐weather events is one of the most secure predictions associated with contemporary climate change, effects of such events on distribution and abundance of climate‐sensitive species remain poorly understood. Montane ecosystems may be especially sensitive to extreme weather because of complex abiotic and biotic interactions that propagate from climate‐driven reductions in snowpack. Snowpack not only protects subnivean biotas from extreme cold, but also influences forage availability through timing of melt‐off and water availability. We related relative abundances of an alpine mammal, the American pika ( Ochotona princeps ), to measures of weather and snowpack dynamics over an 8‐yr period that included before and after a year of record‐low snowpack in Washington, USA. We sought to (1) quantify any change in pika abundance associated with the snowpack anomaly and (2) identify aspects of weather and snowpack that influenced abundance of pikas. Pikas showed a 1‐yr lag response to the snowpack anomaly and exhibited marked declines in abundance at elevations below 1,400 m simultaneous with increased abundances at higher elevations. Atmospheric moisture, indexed by vapor pressure deficit (VPD), was especially important, evidenced by strong support for the top‐ranked model that included the interaction of VPD with snowpack duration. Notably, our novel application of VPD from gridded climate data for analyses of animal abundances shows strong potential for improving species distribution models because VPD represents an important aspect of weather that influences the physiology and habitat of biota. Pikas were apparently affected by cold stress without snowpack at mid elevations, whereas changes to forage associated with snowpack and VPD were influential at high and low elevations. Our results reveal context dependency in pika responses to weather and illustrate how snow drought can lead to rapid change in the abundance of subnivean animals.

Washington↗

Modeling spatiotemporal abundance and movement dynamics using an integrated spatial capture–recapture movement model

Animal movement is a fundamental ecological process affecting the survival and reproduction of individuals, the structure of populations, and the dynamics of communities. Methods to quantify animal movement and spatiotemporal abundances, however, are generally separate and therefore omit linkages between individual-level and population-level processes. We describe an integrated spatial capture–recapture (SCR) movement model to jointly estimate (1) the number and distribution of individuals in a defined spatial region and (2) movement of those individuals through time. We applied our model to a study of polar bears ( Ursus maritimus ) in a 28,125 km 2 survey area of the eastern Chukchi Sea, USA in 2015 that incorporated capture–recapture and telemetry data. In simulation studies, the model provided unbiased estimates of movement, abundance, and detection parameters using a bivariate normal random walk and correlated random walk movement process. Our case study provided detailed evidence of directional movement persistence for both male and female bears, where individuals regularly traversed areas larger than the survey area during the 36-day study period. Scaling from individual- to population-level inferences, we found that densities varied from <0.75 bears/625 km 2 grid cell/day in nearshore cells to 1.6–2.5 bears/grid cell/day for cells surrounded by sea ice. Daily abundance estimates ranged from 53 to 69 bears, with no trend across days. The cumulative number of unique bears that used the survey area increased through time due to movements into and out of the area, resulting in an estimated 171 individuals using the survey area during the study (95% credible interval 124–250). Abundance estimates were similar to a previous multiyear integrated population model using capture–recapture and telemetry data (2008–2016; Regehr et al., Scientific Reports 8:16780, 2018). Overall, the SCR–movement model successfully quantified both individual- and population-level space use, including the effects of landscape characteristics on movement, abundance, and detection, while linking the movement and abundance processes to directly estimate density within a prescribed spatial region and temporal period. Integrated SCR–movement models provide a generalizable approach to incorporate greater movement realism into population dynamics and link movement to emergent properties including spatiotemporal densities and abundances.

eastern Chukchi Sea↗

Weather influences survival probability in two coexisting mammals directly and indirectly via competitive asymmetry

Ecologists have studied the role of interspecific competition in structuring ecological communities for decades. Differential weather effects on animal competitors may be a particularly important factor contributing to the outcome of competitive interactions, though few studies have tested this hypothesis in free-ranging animals. Specifically, weather might influence competitive dynamics by altering competitor densities and/or per-capita competitive effects on demographic vital rates. We used a 9-year data set of marked individuals to test for direct and interactive effects of weather and competitor density on survival probability in two coexisting mammalian congeners: Columbian ground squirrels ( Urocitellus columbianus ) and northern Idaho ground squirrels ( Urocitellus brunneus ). Ambient temperature and precipitation influenced survival probability in both species, but the effects of weather differed between the two species. Moreover, density of the larger Columbian ground squirrel negatively impacted survival probability in the smaller northern Idaho ground squirrel (but not vice versa), and the strength of the negative effect was exacerbated by precipitation. That is, cooler, wetter conditions benefited the larger competitor to the detriment of the smaller species. Our results suggest weather-driven environmental variation influences the competitive equilibrium between ecologically similar mammals of differential body size. Whether future climate change leads to the competitive exclusion of either species will likely depend on the mechanism(s) explaining the coexistence of these competing species. Divergent body size and, hence, differences in thermal tolerance and giving up densities offer potential explanations for the weather-dependent competitive asymmetry we documented, especially if the larger species competitively excludes the smaller species from habitat patches of shared preference via interference.

Idaho↗

Large scale wildlife monitoring studies: Statistical methods for design and analysis

Techniques for estimation of absolute abundance of wildlife populations have received a lot of attention in recent years. The statistical research has been focused on intensive small-scale studies. Recently, however, wildlife biologists have desired to study populations of animals at very large scales for monitoring purposes. Population indices are widely used in these extensive monitoring programs because they are inexpensive compared to estimates of absolute abundance. A crucial underlying assumption is that the population index ( C ) is directly proportional to the population density ( D ). The proportionality constant, β , is simply the probability of 'detection' for animals in the survey. As spatial and temporal comparisons of indices are crucial, it is necessary to also assume that the probability of detection is constant over space and time. Biologists intuitively recognize this when they design rigid protocols for the studies where the indices are collected. Unfortunately, however in many field studios the assumption is clearly invalid. We believe that the estimation of detection probability should be built into the monitoring design through a double sampling approach. A large sample of points provides an abundance index, and a smaller sub-sample of the same points is used to estimate detection probability. There is an important need for statistical research on the design and analysis of these complex studies. Some basic concepts based on actual avian, amphibian, and fish monitoring studies are presented in this article.

Environmetrics↗

Observed and predicted reproduction of Ceriodaphnia dubia exposed to chloride, sulfate, and bicarbonate

Chronic toxicities of Cl - , SO 4 2- , and HCO 3 - to Ceriodaphnia dubia were evaluated in low- and moderate-hardness waters using a three-brood reproduction test method. Toxicity tests of anion mixtures were used to determine interaction effects and to produce models predicting C. dubia reproduction. Effluents diluted with low- and moderate-hardness waters were tested with animals acclimated to low- and moderate-hardness conditions to evaluate the models and to assess the effects of hardness and acclimation. Sulfate was significantly less toxic than Cl - and HCO 3 - in both types of water. Chloride and HCO 3 - toxicities were similar in low-hardness water, but HCO 3 - was the most toxic in moderate-hardness water. Low acute-to-chronic ratios indicate that toxicities of these anions will decrease quickly with dilution. Hardness significantly reduced Cl - and SO 4 2- toxicity but had little effect on HCO 3 - . Chloride toxicity decreased with an increase in Na + concentration, and CO 3 - toxicity may have been reduced by the dissolved organic carbon in effluent. Multivariate models using measured anion concentrations in effluents with low to moderate hardness levels provided fairly accurate predictions of reproduction. Determinations of toxicity for several effluents differed significantly depending on the hardness of the dilution water and the hardness of the water used to culture test animals. These results can be used to predict the contribution of elevated anion concentrations to the chronic toxicity of effluents; to identify effluents that are toxic due to contaminants other than Cl - , SO 4 2- , and HCO 3 - ; and to provide a basis for chemical substitutions in manufacturing processes.

Environmental Toxicology and Chemistry↗

Alternative approaches to vertebrate ecotoxicity tests in the 21st century: A review of developments over the last 2 decades and current status

The need for alternative approaches to the use of vertebrate animals for hazard assessment of chemicals and pollutants has become of increasing importance. It is now the first consideration when initiating a vertebrate ecotoxicity test, to ensure that unnecessary use of vertebrate organisms is minimized wherever possible. For some regulatory purposes, the use of vertebrate organisms for environmental risk assessments has been banned; in other situations, the number of organisms tested has been dramatically reduced or the severity of the procedure refined. However, there is still a long way to go to achieve a complete replacement of vertebrate organisms to generate environmental hazard data. The development of animal alternatives is based not just on ethical considerations but also on reducing the cost of performing vertebrate ecotoxicity tests and in some cases on providing better information aimed at improving environmental risk assessments. The present Focus article provides an overview of the considerable advances that have been made toward alternative approaches for ecotoxicity assessments over the last few decades.

Environmental Toxicology and Chemistry↗

Metal bioavailability models: Current status, lessons learned, considerations for regulatory use, and the path forward

Since the early 2000s, biotic ligand models and related constructs have been a dominant paradigm for risk assessment of aqueous metals in the environment. We critically review 1) the evidence for the mechanistic approach underlying metal bioavailability models; 2) considerations for the use and refinement of bioavailability-based toxicity models; 3) considerations for the incorporation of metal bioavailability models into environmental quality standards; and 4) some consensus recommendations for developing or applying metal bioavailability models. We note that models developed to date have been particularly challenged to accurately incorporate pH effects because they are unique with multiple possible mechanisms. As such, we doubt it is ever appropriate to lump algae/plant and animal bioavailability models; however, it is often reasonable to lump bioavailability models for animals, although aquatic insects may be an exception. Other recommendations include that data generated for model development should consider equilibrium conditions in exposure designs, including food items in combined waterborne–dietary matched chronic exposures. Some potentially important toxicity-modifying factors are currently not represented in bioavailability models and have received insufficient attention in toxicity testing. Temperature is probably of foremost importance; phosphate is likely important in plant and algae models. Acclimation may result in predictions that err on the side of protection. Striking a balance between comprehensive, mechanistically sound models and simplified approaches is a challenge. If empirical bioavailability tools such as multiple-linear regression models and look-up tables are employed in criteria, they should always be informed qualitatively and quantitatively by mechanistic models. If bioavailability models are to be used in environmental regulation, ongoing support and availability for use of the models in the public domain are essential.

Environmental Toxicology and Chemistry↗

Air pollution and wildlife toxicology: An overlooked problem

Since the 1880s, there have been repeated reports worldwide of toxic effects to wildlife from air pollution . Incidents in North America were recorded as early as the 1920s and as recently as last year. The effects have ranged from death and injury to increased incidence of infectious diseases, and they are the result of exposure to both gaseous and particulate emissions. Compared with other threats to wildlife , for instance, pesticides, the toxicological relationship between air pollution and effects in wildlife is not well understood. Our limited understanding is based primarily on reports of symptoms observed in the field and on information extrapolated from studies in livestock and laboratory animals. Few controlled wildlife studies, such as those that have evaluated pesticide effects, have been conducted. Current air quality standards cannot be assumed to protect wildlife from the effects of air pollution . This article reviews the current state of knowledge of air pollution and wildlife toxicology and the continuing threat air pollution presents to wildlife . The available information on reported effects of gaseous and particulate pollutants (i.e., sulfur dioxide, hydrogen sulfide, oxidants, arsenic, cadmium, fluoride, lead and selenium) on terrestrial wildlife , along with similar toxicological data for domestic animals, is summarized. Information on the toxic effects, tolerance levels, pathways of contamination, and risks to wildlife from air pollution is given, and gaps in knowledge are pointed out. Areas where research is needed are identified. Copyright © 1988 SETAC

Environmental Toxicology and Chemistry↗

A resource conservative procedure for comparison of dose-response relationships

The evaluation of effects of toxicants on a wildlife community can be complicated by varying responses among the community's constituent populations. Even within populations, considerable variability in dose-response relations may result from different avenues of exposure to the toxicant. Full-scale investigations of the dose-response relations among a variety of species and avenues of exposure can therefore be prohibitively expensive, whether this expense is measured by the number of experimental animals needed, by the human resources committed to the study, or by laboratory expenses. We propose an abbreviated protocol for investigations of multiple dose-response relations that is designed to limit these expenses. The protocol begins with the judicious choice of a baseline dose-response relation to be estimated by a full-scale study involving a minimum of five doses levels, with 10 subjects per dose level. This relation is then used as the basis for rapid screening of subsequent dose-response relations, which are compared to the baseline relation by testing for differences in the median effective dosages. These secondary studies can consist of as few as 14 animals exposed to the estimated median lethal concentration from the baseline study. We describe MS-DOS-compatible software available from the authors that can be used to analyze these data.

Environmental Toxicology and Chemistry↗

Nutrient transport to the Swan - Canning Estuary, Western Australia

Catchment nutrient availability in Western Australia is primarily controlled by the disposal of animal waste and the type and rate of fertilizer application, particularly on the relatively narrow (~25 km wide), sandy coastal plain. Nitrogen (N) and phosphorous (P) concentrations and fluxes during the wet season of 15 tributaries, including four urban drains to the Swan-Canning Estuary, were evaluated from 1986 to 1992 and additionally concentrations only were evaluated throughout the year from 1993 to 1996. Concentrations of filtered reactive P (FRP) and total P (TP) were generally low, with the volume-weighted means for all sites being 0.06 mg 1-1 and 0.12 mg 1-1 respectively. The urban drains had higher TP concentrations (volume-weighted mean of 0.21 mg 1-1) than the streams (0.12 mg 1-1), with the high concentrations associated with particulate matter. Total inorganic N (TIN, NH4N plus NO3N) and total N (TN), which is of interest to eutrophic status of the N-limited estuary, were likewise low, compared with other developed areas having a similar climate. Both TIN and TN were higher in the urban drains (0.76 mg 1-1 and 1.5 mg 1-1 respectively) than the streams (0.31 mg 1-1 and 1.2 mg 1-1 respectively). The Avon River, which drains 98.5% of the 121 000 km2 catchment area, contributes most of the N (0.03 kg ha-1 year-1 or 65%) and a high percentage of the P (<0.01 kg ha-1 year-1 or 32%) to the estuaries. The Avon River nutrient fluxes are much less than other tributaries closer to the estuary. The coastal plain receives significantly higher rainfall (1,200 mm year-1) and has more intense horticulture and animal production than inland areas (<300 mm year-1). Annual rainfall is seasonal, occuring primarily from May through December. The surficial aquifers on the coastal plain generally are sandy with a low nutrient retention capacity, and rapidly transmit soluble and colloidal material in subsurface flow. Ellen Brook, on the coastal plain, drains pastures treated with superphosphate and has the highest FRP (0.51 mg 1-1), TP (0.7 mg 1-1) and TN (2.1 mg 1-1) of any tributary to the estuary. The coastal plain is also undergoing urbanization, particularly in areas adjacent to the estuary. Nutrients are subsequently available for transport during the onset of seasonal wet weather. Perennial baseflow from urban areas is an important source of nutrients. Water yield from the urban areas was high, being as much as 50% of annual rainfall. The timing of the nutrients delivered by the tributaries may be an important control on estuarine ecology. Copyright ?? 2001 John Wiley and Sons, Ltd.

Hydrological Processes↗

Wildlife ecological risk assessment in the 21st century: Promising technologies to assess toxicological effects

Despite advances in toxicity testing and development of new approach methodologies (NAMs) for hazard assessment, the ecological risk assessment (ERA) framework for terrestrial wildlife (i.e., air-breathing amphibians, reptiles, birds, and mammals) has remained unchanged for decades. While survival, growth, and reproductive endpoints derived from whole animal toxicity tests are central to hazard assessment, non-standard measures of biological effects at multiple levels of biological organization (e.g., molecular, cellular, tissue, organ, organism, population, community, ecosystem) have potential to enhance the relevance of prospective and retrospective wildlife ERAs. Other factors (e.g., indirect effects of contaminants on food supplies and infectious disease processes) are influenced by toxicants at individual, population, and community levels, and need to be factored into chemically-based risk assessments to enhance the “eco” component of ERAs. Regulatory and logistical challenges often relegate such non-standard endpoints and indirect effects to post-registration evaluations of pesticides and industrial chemicals, and contaminated site evaluations. While NAMs are being developed, to date their applications in ERAs focused on wildlife have been limited. No single magic tool or model will address all uncertainties in hazard assessment. Modernizing wildlife ERAs will likely entail combinations of laboratory and field-derived data at multiple levels of biological organization, knowledge collection solutions (e.g., systematic review, adverse outcome pathway frameworks), and inferential methods that facilitate integrations and risk estimations focused on species, populations, interspecific extrapolations, and ecosystem services modeling, with less dependence on whole animal data and simple hazard ratios.

Integrated Environmental Assessment and Management↗

Demographic and temporal variations in immunity and condition of polar bears (Ursus maritimus) from the southern Beaufort Sea

Assessing the health and condition of animals in their natural environment can be problematic. Many physiological metrics, including immunity, are highly influenced by specific context and recent events to which researchers may be unaware. Thus, using a multifaceted physiological approach and a context-specific analysis encompassing multiple time scales can be highly informative. Ecoimmunological tools in particular can provide important indications to the health of animals in the wild. We collected blood and hair samples from free-ranging polar bears (Ursus maritimus) in the southern Beaufort Sea and examined the influence of sex, age, and reproductive status on metrics of immunity, stress, and body condition during 2013–2015. We examined metrics of innate immunity (bactericidal ability and lysis) and stress (hair cortisol, reactive oxygen species, and oxidative barrier), in relation to indices of body condition considered to be short term (urea to creatinine ratio; UC ratio) and long term (storage energy and body mass index). We found the factors of sex, age, and reproductive status of the bear were critical for interpreting different physiological metrics. Additionally, the metrics of body condition were important predictors for stress indicators. Finally, many of these metrics differed between years, illustrating the need to examine populations on a longer time scale. Taken together, this study demonstrates the complex relationship between multiple facets of physiology and how interpretation requires us to examine individuals within a specific context.

Alaska↗

Sympatric cattle grazing and desert bighorn sheep foraging

Foraging behavior affects animal fitness and is largely dictated by the resources available to an animal. Understanding factors that affect forage resources is important for conservation and management of wildlife. Cattle sympatry is proposed to limit desert bighorn population performance, but few studies have quantified the effect of cattle foraging on bighorn forage resources or foraging behavior by desert bighorn. We estimated forage biomass for desert bighorn sheep in 2 mountain ranges: the cattle-grazed Caballo Mountains and the ungrazed San Andres Mountains, New Mexico. We recorded foraging bout efficiency of adult females by recording feeding time/step while foraging, and activity budgets of 3 age-sex classes (i.e., adult males, adult females, yearlings). We also estimated forage biomass at sites where bighorn were observed foraging. We expected lower forage biomass in the cattle-grazed Caballo range than in the ungrazed San Andres range and lower biomass at cattle-accessible versus inaccessible areas within the Caballo range. We predicted bighorn would be less efficient foragers in the Caballo range. Groundcover forage biomass was low in both ranges throughout the study (Jun 2012&ndash;Nov 2013). Browse biomass, however, was 4.7 times lower in the Caballo range versus the San Andres range. Bighorn in the Caballo range exhibited greater overall daily travel time, presumably to locate areas of higher forage abundance. By selecting areas with greater forage abundance, adult females in the Caballo range exhibited foraging bout efficiency similar to their San Andres counterparts but lower overall daily browsing time. We did not find a significant reduction in forage biomass at cattle-accessible areas in the Caballo range. Only the most rugged areas in the Caballo range had abundant forage, potentially a result of intensive historical livestock use in less rugged areas. Forage conditions in the Caballo range apparently force bighorn to increase foraging effort by feeding only in areas where adequate forage remains.

New Mexico↗

A broader definition of occupancy: A reply to Hayes and Monofils

Occupancy models are widely used to analyze presence&ndash;absence data for a variety of taxa while accounting for observation error (MacKenzie et al. 2002, 2006; Tyre et al. 2003; Royle and Dorazio 2008). Hayes and Monfils (2015) question their use for analyzing avian point count data based on purported violations of model assumptions incurred by avian mobility. Animal mobility is an important consideration, not just for occupancy models, but for a variety of population and habitat models (Boyce 2006, Royle et al. 2009, Manning and Goldberg 2010, Dormann et al. 2013, Renner et al. 2015). Nevertheless, we believe the ultimate conclusions of Hayes and Monfils are shortsighted mainly due to a narrow interpretation of occupancy. Rather than turn away from the use of occupancy models, we believe they remain an appropriate method for analyzing many data sets collected from avian point count surveys. Further, we suggest that there is value in having a broader and more nuanced interpretation of occupancy that incorporates the potential for animal movement.

Journal of Wildlife Management↗

Population ecology of Roosevelt elk: Conservation and management in Redwood National and State Parks. Butch Weckerly. 2017. University of Nevada Press, Reno, Nevada, USA. 224 pp. $54.95 hardback. ISBN 978- 1943859504.

Long-term research on large ungulate populations typically conjures perceptions of extensive (and expensive) animal capture and telemetry work, and subsequent advanced modeling of resource selection and population dynamics that inform management decisions. In contrast, studies lacking a telemetry component are often limited to animal behavior or natural history. Although compelling from a standpoint of advancing understanding of ecological and evolutionary processes, results from the latter can be unfairly labeled as esoteric because they are not easily transferable to resource managers or may not provide exceptional interest to a general public drawn to these charismatic megafuana. Such dichotomies are not predetermined, however, because study conditions exist where dedicated academic researchers on tight budgets can achieve results relevant to ecology and management.

Journal of Wildlife Management↗

Winter versus summer habitat selection in a threatened ground squirrel

Hibernation is a strategy many species employ to survive periods of thermal stress or resource shortage (e.g., harsh thermal conditions, food limitations) and habitat requirements of hibernating species may differ between summer (the active season) and winter (during hibernation). Accounting for seasonal differences in habitat affinities will help ensure that management actions are more beneficial and land-use policies are more appropriate. The northern Idaho ground squirrel ( Urocitellus brunneus ) is a federally listed threatened species that is in decline and hibernates for approximately 8 months per year. We collared northern Idaho ground squirrels in Adams County, Idaho from 2013–2017. The majority of northern Idaho ground squirrels we collared selected hibernacula outside of the areas they used during the active season. Furthermore, habitat features of hibernacula locations differed from habitat features of active-season areas. Hibernacula locations had greater canopy closure compared to active-season locations (36.9% and 7.0% canopy closure, respectively) and hibernaculum habitat features (particularly distance to nearest log) influenced overwinter survival. Our results suggest that recovery efforts for northern Idaho ground squirrels should include protection and management for the full range of habitat conditions used throughout summer and winter. More broadly, we emphasize the need to identify and protect habitat during all seasons because habitat requirements can differ substantially during different portions of an animal's annual cycle and effective conservation will require management of year-round habitat needs.

Idaho↗