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Camera traps and mark-resight models: The value of ancillary data for evaluating assumptions

Unbiased estimators of abundance and density are fundamental to the study of animal ecology and critical for making sound management decisions. Capture–recapture models are generally considered the most robust approach for estimating these parameters but rely on a number of assumptions that are often violated but rarely validated. Mark-resight models, a form of capture–recapture, are well suited for use with noninvasive sampling methods and allow for a number of assumptions to be relaxed. We used ancillary data from continuous video and radio telemetry to evaluate the assumptions of mark-resight models for abundance estimation on a barrier island raccoon (Procyon lotor) population using camera traps. Our island study site was geographically closed, allowing us to estimate real survival and in situ recruitment in addition to population size. We found several sources of bias due to heterogeneity of capture probabilities in our study, including camera placement, animal movement, island physiography, and animal behavior. Almost all sources of heterogeneity could be accounted for using the sophisticated mark-resight models developed by McClintock et al. (2009b) and this model generated estimates similar to a spatially explicit mark-resight model previously developed for this population during our study. Spatially explicit capture–recapture models have become an important tool in ecology and confer a number of advantages; however, non-spatial models that account for inherent individual heterogeneity may perform nearly as well, especially where immigration and emigration are limited. Non-spatial models are computationally less demanding, do not make implicit assumptions related to the isotropy of home ranges, and can provide insights with respect to the biological traits of the local population.

North Carolina

Moderate susceptibility to subcutaneous plague (Yersinia pestis) challenge in vaccine-treated and untreated Sonoran deer mice (Peromyscus maniculatus sonoriensis) and northern grasshopper mice (Onychomys leucogaster)

The variable response of wild mice to Yersinia pestis infection, the causative agent of plague, has generated much speculation concerning their role in the ecology of this potentially lethal disease. Researchers have questioned the means by which Y. pestis is maintained in nature and also sought methods for managing the disease. Here we assessed the efficacy of a new tool, the sylvatic plague vaccine (SPV), in wild-caught northern grasshopper mice ( Onychomys leucogaster ) and commercially acquired Sonoran deer mice ( Peromyscus maniculatus sonoriensis ). More than 40% of the animals survived a subcutaneous Y. pestis challenge of 175,000 colony forming units (over 30,000 times the white mouse 50% lethal dose) in both vaccine-treated and control groups. Our results indicate that SPV distribution is unlikely to protect adult mice from plague infection in field settings and corroborate the heterogeneous response to Y. pestis infection in mice reported by others.

Journal of Wildlife Diseases

Basis function models for animal movement

Advances in satellite-based data collection techniques have served as a catalyst for new statistical methodology to analyze these data. In wildlife ecological studies, satellite-based data and methodology have provided a wealth of information about animal space use and the investigation of individual-based animal–environment relationships. With the technology for data collection improving dramatically over time, we are left with massive archives of historical animal telemetry data of varying quality. While many contemporary statistical approaches for inferring movement behavior are specified in discrete time, we develop a flexible continuous-time stochastic integral equation framework that is amenable to reduced-rank second-order covariance parameterizations. We demonstrate how the associated first-order basis functions can be constructed to mimic behavioral characteristics in realistic trajectory processes using telemetry data from mule deer and mountain lion individuals in western North America. Our approach is parallelizable and provides inference for heterogenous trajectories using nonstationary spatial modeling techniques that are feasible for large telemetry datasets. Supplementary materials for this article are available online.

Journal of the American Statistical Association

Feeding Ecology of Breeding Blue-Winged Teals

A 5-year investigation of factors influencing the selection of foods consumed by blue-winged teals ( Anas discors ) during the breeding season in the glaciated prairie region of south-central North Dakota showed that birds first arriving on the breeding grounds consumed a diet consisting of 45 percent invertebrates. The proportion of animal foods in the diet increased to 95 percent at the onset of the nesting season. The quality and quantity of foods selected at any given time were influenced by the biological demands and morphological adaptations of the bird, the behavior and ecology of the invertebrates selected as foods, and the general nature of the aquatic ecosystems as determined by the hydrology and geology of the area and modified by land use and weather. Feeding activities changed significantly when food availability within the aquatic ecosystem changed. During the spring and early summer, temporary and seasonal wetlands, if not severely disturbed, were of paramount importance to breeding blue-winged teals since they provided abundant and readily available, high protein, animal foods. Later in the summer when seasonal wetlands began to dry up, insects began to emerge in the semipermanent ponds and lakes, and feeding intensity shifted to these more permanent waters. This trend, however, was often reversed temporarily during the early summer following heavy precipitation that refilled shallow water areas and again stimulated invertebrate development.

North Dakota

Remote ecological monitoring with smartphones and tasker

Researchers have increasingly used autonomous monitoring units to record animal sounds, track phenology with timed photographs, and snap images when triggered by motion. We piloted the use of smartphones to monitor wildlife in the Riverside East Solar Energy Zone (California) and at Indiana Dunes National Park (Indiana). For both efforts, we established remote autonomous monitoring stations in which we housed an Android smartphone in a weather-proof box mounted to a pole and powered by solar panels. We connected each smartphone to a Google account, and the smartphone received its recording/photo schedule daily via a Google Calendar connection when in data transmission mode. Phones were automated by Tasker, an Android application for automating cell phone tasks. We describe a simple approach that could be adopted by others who wish to use nonproprietary methods of data collection and analysis.

California, Indiana

Integrating socioecological suitability with human-wildlife conflict risk: Case study for translocation of a large ungulate

Translocations are essential for re-establishing wildlife populations. As they sometimes fail, it is critical to assess factors that influence their success pre-translocation. Socioecological suitability models (SESMs) integrate social acceptance and ecological suitability to enable identification of areas where wildlife populations will expand, which makes it likely that SESMs will also be useful for predicting translocation success. To inform site selection for potential elk Cervus canadensis reintroduction to north-eastern Minnesota, United States, we developed broadscale maps of social acceptance from surveys of local residents and landowners, animal use equivalence (AUE) from forage measured in the field and empirical conflict risk from geospatial data. Resulting SESMs integrated social acceptance favourability scores, AUE and conflict risk, and weighted SESMs showed the relative influences of acceptance and conflict. Social acceptance was positive for local residents and landowners (mean ≥ 5.4; scale of 1–7). AUE (scaled to an elk home range) ranged between 1 and 9 elk/16 km 2 during winter, and from 14 to 83 elk/16 km 2 during summer. Human–elk conflict risk was low (mean ≤ 0.10; scaled 0–1), increasing from north to south. Geographical distributions differed for social acceptance, AUE and conflict risk, and weighted SESMs revealed unsuitable areas that were otherwise obscured. Synthesis and applications . Integrating human–wildlife conflict risk into SESMs shows where social acceptance of translocated species is likely to erode, even where viewed favourably pre-translocation, to inform translocation planning by highlighting interactions between key factors. Such integrated models supplement existing reintroduction biology frameworks by supporting decision-making and knowledge development. In north-eastern Minnesota, natural resource managers who are considering elk reintroductions are using SESMs reported here to identify where human–elk conflict is unlikely to result in an isolated elk population and where addressing concerns for area residents about conflict risk is essential.

Minnesota

Harvesting wildlife affected by climate change: a modelling and management approach for polar bears

The conservation of many wildlife species requires understanding the demographic effects of climate change, including interactions between climate change and harvest, which can provide cultural, nutritional or economic value to humans. We present a demographic model that is based on the polar bear Ursus maritimus life cycle and includes density-dependent relationships linking vital rates to environmental carrying capacity ( K ). Using this model, we develop a state-dependent management framework to calculate a harvest level that (i) maintains a population above its maximum net productivity level (MNPL; the population size that produces the greatest net increment in abundance) relative to a changing K , and (ii) has a limited negative effect on population persistence. Our density-dependent relationships suggest that MNPL for polar bears occurs at approximately 0·69 (95% CI = 0·63–0·74) of K . Population growth rate at MNPL was approximately 0·82 (95% CI = 0·79–0·84) of the maximum intrinsic growth rate, suggesting relatively strong compensation for human-caused mortality. Our findings indicate that it is possible to minimize the demographic risks of harvest under climate change, including the risk that harvest will accelerate population declines driven by loss of the polar bear's sea-ice habitat. This requires that (i) the harvest rate – which could be 0 in some situations – accounts for a population's intrinsic growth rate, (ii) the harvest rate accounts for the quality of population data (e.g. lower harvest when uncertainty is large), and (iii) the harvest level is obtained by multiplying the harvest rate by an updated estimate of population size. Environmental variability, the sex and age of removed animals and risk tolerance can also affect the harvest rate. Synthesis and applications . We present a coupled modelling and management approach for wildlife that accounts for climate change and can be used to balance trade-offs among multiple conservation goals. In our example application to polar bears experiencing sea-ice loss, the goals are to maintain population viability while providing continued opportunities for subsistence harvest. Our approach may be relevant to other species for which near-term management is focused on human factors that directly influence population dynamics within the broader context of climate-induced habitat degradation.

Journal of Applied Ecology

Experimentally derived salinity tolerance of hatchling Burmese pythons (Python molurus bivittatus) from the Everglades, Florida (USA)

In a laboratory setting, we tested the ability of 24 non-native, wild-caught hatchling Burmese pythons (Python molurus bivittatus) collected in the Florida Everglades to survive when given water containing salt to drink. After a one-month acclimation period in the laboratory, we grouped snakes into three treatments, giving them access to water that was fresh (salinity of 0, control), brackish (salinity of 10), or full-strength sea water (salinity of 35). Hatchlings survived about one month at the highest marine salinity and about five months at the brackish-water salinity; no control animals perished during the experiment. These results are indicative of a "worst-case scenario", as in the laboratory we denied access to alternate fresh-water sources that may be accessible in the wild (e.g., through rainfall). Therefore, our results may underestimate the potential of hatchling pythons to persist in saline habitats in the wild. Because of the effect of different salinity regimes on survival, predictions of ultimate geographic expansion by non-native Burmese pythons that consider salt water as barriers to dispersal for pythons may warrant re-evaluation, especially under global climate change and associated sea-level-rise scenarios.

Florida

Drivers of survival of translocated tortoises

Translocation of animals, especially for threatened and endangered species, is a currently popular but very challenging activity. We translocated 158 adult Agassiz's desert tortoises ( Gopherus agassizii ), a threatened species, from the National Training Center, Fort Irwin, in the central Mojave Desert in California, USA, to 4 plots as part of a long-distance, hard-release, mitigation-driven translocation to prevent deaths from planned military maneuvers. We monitored demographic and behavioral variables of tortoises fitted with radio-transmitters from 2008 to 2018. By the end of the project, 17.72% of tortoises were alive, 65.82% were dead, 15.19% were missing, and 1.27% were removed from the study because they returned to Fort Irwin. Mortality was high during the first 3 years: >50% of the released animals died, primarily from predation. Thereafter, mortality declined but remained high. After 10.5 years, survival was highest, 37.50% (15/40), on the plot closest to original home sites, whereas from 2.56% to 23.68% remained alive on the other 3 release plots. Surviving tortoises settled early, repeatedly using locations where they constructed burrows, compared with tortoises that died or disappeared. Models of behavioral and other variables indicated that numbers of repeatedly used locations (burrows) were a driver of survival throughout the study, although plot location, size and sex of tortoises, and distance traveled were contributors, especially during early years. Because >50% mortality occurred, we considered this translocation unsuccessful. The study area appeared to be an ecological sink with historical and current anthropogenic uses contributing to habitat degradation and a decline in both the resident and released tortoises. Our findings will benefit design and selection of future translocation areas.

California

Mortality of Siberian polecats and black-footed ferrets released onto prairie dog colonies

Black-footed ferrets ( Mustela nigripes ) likely were extirpated from the wild in 1985–1986, and their repatriation depends on captive breeding and reintroduction. Postrelease survival of animals can be affected by behavioral changes induced by captivity. We released neutered Siberian polecats ( M. eversmanii ), close relatives of ferrets, in 1989–1990 on black-tailed prairie dog ( Cynomys ludovicianus ) colonies in Colorado and Wyoming initially to test rearing and reintroduction techniques. Captive-born polecats were reared in cages or cages plus outdoor pens, released from elevated cages or into burrows, and supplementally fed or not fed. We also translocated wild-born polecats from China in 1990 and released captive-born, cage-reared black-footed ferrets in 1991, the 1st such reintroduction of black-footed ferrets. We documented mortality for 55 of 92 radiotagged animals in these studies, mostly due to predation (46 cases). Coyotes ( Canis latrans ) killed 31 ferrets and polecats. Supplementally fed polecats survived longer than nonprovisioned polecats. With a model based on deaths per distance moved, survival was highest for wild-born polecats, followed by pen-experienced, then cage-reared groups. Indexes of abundance (from spotlight surveys) for several predators were correlated with mortality rates of polecats and ferrets due to those predators. Released black-footed ferrets had lower survival rates than their ancestral population in Wyoming, and lower survival than wild-born and translocated polecats, emphasizing the influence of captivity. Captive-born polecats lost body mass more rapidly postrelease than did captive-born ferrets. Differences in hunting efficiency and prey selection provide further evidence that these polecats and ferrets are not ecological equivalents in the strict sense.

Journal of Mammalogy

Space between houses influences movement and habitat selection of ungulates: Width as a novel metric of development

Wildlife often lose access to habitat due to housing development. The magnitude of indirect habitat loss can be conditional on the configuration of individual houses, but commonly used metrics (i.e. density or distance) can overlook the configuration of development. We introduce a novel framework to index the configuration of development based on the width of space between houses and associated structures. We use resource selection functions to assess the degree that GPS-collared elk ( Cervus canadensis ), mule deer ( Odocoileus hemionus ), pronghorn and moose ( Alces alces ) on winter range and on migration routes use space between houses within northwest Wyoming, USA, near the towns of Cody and Jackson. Further, to help inform regulations aiming to promote wildlife movement across a gradient of land uses, we differentiated between individuals residing in primarily rural and exurban areas. Rural populations of elk, mule deer, pronghorn and moose avoided spaces narrower than 2 km and never used spaces narrower than 50 m between houses, whereas exurban populations of elk, mule deer and moose selected for spaces narrower than 2 km but avoided spaces narrower than 50 m. We identified cutoffs in rural and exurban areas where space may become too narrow for most animals to use. Through this metric, managers and policy makers can inform the necessary width to maintain wildlife movement through corridors. Our width metric can be applied to other systems and our workflow is publicly available ( https://wildlifemovetools.org/width-calculator ) so users can estimate the width of space between structures in their focal areas. Synthesis and applications . In rural areas, maintaining spaces between houses >500 m will likely facilitate ungulate movement. To fully conserve functional habitat, such as unimpeded habitat use along migration corridors, maintaining spaces approximately >2.5 km between houses will likely be necessary.

Wyoming

Feeding ecology of northern pintails and green-winged teal wintering in California

The feeding ecology of northern pintails (Anas acuta) and green-winged teal (A. crecca) was examined from October through February 1979-81 in 4 major seasonal marsh types in the Central Valley, California. The esophagi of 262 pintails contained 72.3% plant seeds and 27.7% animal matter. The esophagi of 173 green-winged teal contained 62.3% plant seeds and 37.6% animal matter. Swamp timothy (Heleochloa schoenoides) caryopses, chironomid midge larvae, and common barnyardgrass (Echinochloa crusgalli) caryopses formed >50% of the diet of both species. Both species were highly opportunistic and generally shifted their food habits seasonally to the most available foods. Animal matter increased seasonally in the diets of both and formed about 60% of the foods eaten during January and February compared to only about 8% in October and 17% in December. Both species used open water marsh habitats almost exclusively in daytime but they used densely vegetated marshes almost exclusively at night. Management recommendations based on the food habits and habitat use patterns of pintails and green-winged teal are offered.

Journal of Wildlife Management

Hierarchical models of animal abundance and occurrence

Much of animal ecology is devoted to studies of abundance and occurrence of species, based on surveys of spatially referenced sample units. These surveys frequently yield sparse counts that are contaminated by imperfect detection, making direct inference about abundance or occurrence based on observational data infeasible. This article describes a flexible hierarchical modeling framework for estimation and inference about animal abundance and occurrence from survey data that are subject to imperfect detection. Within this framework, we specify models of abundance and detectability of animals at the level of the local populations defined by the sample units. Information at the level of the local population is aggregated by specifying models that describe variation in abundance and detection among sites. We describe likelihood-based and Bayesian methods for estimation and inference under the resulting hierarchical model. We provide two examples of the application of hierarchical models to animal survey data, the first based on removal counts of stream fish and the second based on avian quadrat counts. For both examples, we provide a Bayesian analysis of the models using the software WinBUGS.

Journal of Agricultural, Biological, and Environme

Invasive species management restores a plant-pollinator mutualism in Hawaii

1.The management and removal of invasive species may give rise to unanticipated changes in plant–pollinator mutualisms because they can alter the composition and functioning of plant–pollinator interactions in a variety of ways. To utilize a functional approach for invasive species management, we examined the restoration of plant–pollinator mutualisms following the large-scale removal of an invasive nectar thief and arthropod predator, Vespula pensylvanica. 2.We reduced V. pensylvanica populations in large plots managed over multiple years to examine the response of plant–pollinator mutualisms and the fruit production of a functionally important endemic Hawaiian tree species, Metrosideros polymorpha. To integrate knowledge of the invader's behaviour and the plant's mating system, we determined the efficacy of V. pensylvanica as a pollinator of M. polymorpha and quantified the dependence of M. polymorpha on animal pollination (e.g. level of self-compatibility and pollen limitation). 3.The reduction of V. pensylvanica in managed sites, when compared to unmanaged sites, resulted in a significant increase in the visitation rates of effective bee pollinators (e.g. introduced Apis mellifera and native Hylaeus spp.) and in the fruit production of M. polymorpha. 4.Apis mellifera, following the management of V. pensylvanica, appears to be acting as a substitute pollinator for M. polymorpha, replacing extinct or threatened bird and bee species in our study system. 5.Synthesis and applications. Fruit production of the native M. polymorpha was increased after management of the invasive pollinator predator V. pensylvanica; however, the main pollinators were no longer native but introduced. This research thus demonstrates the diverse impacts of introduced species on ecological function and the ambiguous role they play in restoration. We recommend incorporating ecological function and context into invasive species management as this approach may enable conservation managers to simultaneously minimize the negative and maximize the positive impacts (e.g. taxon substitution) of introduced species. Such novel restoration approaches are needed, especially in highly degraded ecosystems.

Hawai'i

Managing more than the mean: Using quantile regression to identify factors related to large elk groups

Summary Animal group size distributions are often right-skewed, whereby most groups are small, but most individuals occur in larger groups that may also disproportionately affect ecology and policy. In this case, examining covariates associated with upper quantiles of the group size distribution could facilitate better understanding and management of large animal groups. We studied wintering elk groups in Wyoming, where group sizes span several orders of magnitude, and issues of disease, predation and property damage are affected by larger group sizes. We used quantile regression to evaluate relationships between the group size distribution and variables of land use, habitat, elk density and wolf abundance to identify conditions important to larger elk groups. We recorded 1263 groups ranging from 1 to 1952 elk and found that across all quantiles of group size, group sizes were larger in open habitat and on private land, but the largest effect occurred between irrigated and non-irrigated land [e.g. the 90th quantile group size increased by 135 elk (95% CI = 42, 227) on irrigation]. Only upper quantile group sizes were positively related to broad-scale measures of elk density and wolf abundance. For wolf abundance, this effect was greater on elk groups found in open habitats and private land than those in closed habitats or public land. If we had limited our analysis to mean or median group sizes, we would not have detected these effects. Synthesis and applications . Our analysis of elk group size distributions using quantile regression suggests that private land, irrigation, open habitat, elk density and wolf abundance can affect large elk group sizes. Thus, to manage larger groups by removal or dispersal of individuals, we recommend incentivizing hunting on private land (particularly if irrigated) during the regular and late hunting seasons, promoting tolerance of wolves on private land (if elk aggregate in these areas to avoid wolves) and creating more winter range and varied habitats. Relationships to the variables of interest also differed by quantile, highlighting the importance of using quantile regression to examine response variables more completely to uncover relationships important to conservation and management.

Wyoming

Human activities and weather drive contact rates of wintering elk

Wildlife aggregation patterns can influence disease transmission. However, limited research evaluates the influence of anthropogenic and natural factors on aggregation. Many managers would like to reduce wildlife contact rates, driven by aggregation, to limit disease transmission. We develop a novel analytical framework to quantify how management activities such as supplemental feeding and hunting versus weather drive contact rates while accounting for correlated contacts. We apply the framework to the National Elk Refuge (NER), Wyoming, USA, where the probable arrival of chronic wasting disease (CWD) has magnified concerns. We used a daily proximity index to measure contact rates among 68 global positioning system collared elk from 2016 to 2019. We modelled contact rates as a function of abiotic weather‐related effects, anthropogenic effects and aggregation from the prior day. The winter of 2017–2018 had greater natural forage availability and little snow, which led to a rare non‐feeding year on the NER and provided a unique opportunity to evaluate the effect of feeding on contact rates relative to other conditions. Supplemental feeding was the strongest predictor of aggregation, and contact rates were 2.6 times larger while feeding occurred compared to the baseline rate (0.34 and 0.13, respectively). Snow‐covered area was the second strongest predictor of contact rates highlighting the importance of abiotic factors to elk aggregation, but this effect had half the strength of feeding. These results are the first to show, even in animals that congregate naturally, how greatly supplemental feeding amplifies aggregation. Contact rates were also 23% lower during times when elk hunting was active (0.10) compared to the baseline. Synthesis and applications . Supplemental feeding increased contacts between elk well above the natural effects of weather, even after accounting for correlated movement expected in wintering ungulates. Similarly, differences in hunting season timing with adjacent areas led to an increase in contacts, suggesting an additional management option for reducing aggregation. The analytical framework presented supports the evaluation of temporally varying management actions that influence aggregation broadly and can be easily implemented whether the interest in changing aggregation is related to reduction of disease transmission, human–wildlife conflict or inter‐species competition.

Wyoming

Using spatial capture–recapture to elucidate population processes and space-use in herpetological studies

The cryptic behavior and ecology of herpetofauna make estimating the impacts of environmental change on demography difficult; yet, the ability to measure demographic relationships is essential for elucidating mechanisms leading to the population declines reported for herpetofauna worldwide. Recently developed spatial capture–recapture (SCR) methods are well suited to standard herpetofauna monitoring approaches. Individually identifying animals and their locations allows accurate estimates of population densities and survival. Spatial capture–recapture methods also allow estimation of parameters describing space-use and movement, which generally are expensive or difficult to obtain using other methods. In this paper, we discuss the basic components of SCR models, the available software for conducting analyses, and the experimental designs based on common herpetological survey methods. We then apply SCR models to Red-backed Salamander ( Plethodon cinereus ), to determine differences in density, survival, dispersal, and space-use between adult male and female salamanders. By highlighting the capabilities of SCR, and its advantages compared to traditional methods, we hope to give herpetologists the resource they need to apply SCR in their own systems.

Journal of Herpetology

Testing assumptions in the use of PIT tags to study movement of Plethodon salamanders

Studying the movements of organisms that live underground for at least a portion of their life history is challenging, given the state of current technology. Passive integrated transponders (PIT tags) provide a way to individually identify and, more recently, study the movement of smaller animals, including those that make subterranean movements. However, there are widespread assumptions of the use of PIT tags that remain problematic. We tested the effects of PIT-tag implantation on growth and survival, along with the effects of electromagnetic fields for reading PIT tags on behavior, of the smallest salamander that has been PIT-tagged: the Red-Backed Salamander. We found no effect of PIT tags on growth or survival. Using a mesocosm experiment, we also found that electromagnetic effects associated with reading PIT tags, had no effect on salamander behavior. Further, we describe a novel PIT antenna and soil mesocosm experimental arena for studying belowground movements of woodland salamanders. Collectively, these studies suggest that the use of PIT tags do not influence the growth, survival, or behavior of Red-Backed Salamanders. Given the challenges of studying salamanders that live underground and the impending changes in climate and landscapes, this research suggests that PIT tags remain a viable tool for studying the movement ecology of salamanders under global change.

Journal of Herpetology