Meet our grantees

Wild Animal Initiative funds academic research on high-priority questions in wild animal welfare.

The goal of our grants program is to fund research that deepens scientific knowledge of the welfare of wild animals in order to better understand how to improve the welfare of as many wild animals as possible, regardless of what causes the threats to their well-being.

We showcase our grantees and their projects here and will be adding more in the coming weeks and months.

Assessing the Welfare of Wild Olive Baboons (Papio anubis) at the Uaso Ngiro Baboon Project, Laikipia, Kenya

Grantee: Monica Wakefield

Institution: Northern Kentucky University and Iowa State University

Project summary

This project aims to assess the welfare of wild baboons using a holistic approach. The study takes advantage of almost 40 years of long-term ecological, demographic, health, and behavioral information on >170 individually known baboons, and will examine how various factors such as age, sex, and social rank correlate with individual welfare measures such as incidence of injury and disease, body mass, and pro-social or agonistic behaviors. The annual scale of the dataset will also enable the researchers to test how events such as droughts have affected welfare during specific periods, and how these effects may have varied according to individual animals’ demographic profiles and biographies.

Grantee: Monica Wakefield

 

Institutions: Northern Kentucky University and Iowa State University, United States

Grant amount: $29,800

 

Grant type: Small grants

Focal species: Wild olive baboon (Papio anubis)

 

Conservation status: Least concern

Disciplines: Animal welfare science, animal behavior, primatology

 

Research location: Kenya, United States


Project summary

This project aims to assess the welfare of wild baboons using a holistic approach. The study takes advantage of almost 40 years of long-term ecological, demographic, health, and behavioral information on >170 individually known baboons, and will examine how various factors such as age, sex, and social rank correlate with individual welfare measures such as incidence of injury and disease, body mass, and pro-social or agonistic behaviors. The annual scale of the dataset will also enable the researchers to test how events such as droughts have affected welfare during specific periods, and how these effects may have varied according to individual animals’ demographic profiles and biographies.

Why we funded this project

We are excited by this project’s analysis of a long-term longitudinal dataset because of the importance of understanding how wild animals’ welfare varies with demographic factors such as age and sex, as these groups often face different challenges and have different ecological and behavioral requirements, and negative welfare impacts that fall on young individuals may also have ripple effects throughout their lives. We also appreciate this project’s holistic approach. The long-term monitoring means it is possible to consider not only the usual downstream welfare indicators based on health and behavior, but also upstream factors that might influence them, such as social interactions and exposure to predators.


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Assessing the anthropogenic impacts, long-term health, and welfare of elasmobranch species within San Francisco Bay, California

Grantee: Meghan Holst

Institution: University of California, Davis

Project summary

The San Francisco Bay Estuary is used by individuals belonging to several elasmobranch populations during critical periods of their life history. But it is also dredged, used as a major harbor for ships, and fished, with sharks and their prey targeted. This project will measure stress physiology and blood contaminants to evaluate whether the health and welfare of elasmobranchs is threatened within San Francisco Bay. It will also use stable isotope analysis to evaluate the dietary needs and sensitivities of San Francisco Bay’s elasmobranch species to determine whether individuals are threatened with hunger due to fishing, and to map relationships within the ecosystem.

Grantee: Meghan Holst

 

Institution: University of California, Davis, United States

Grant amount: $19,200

 

Grant type: Small grants

Focal species: Shark (Selachimorpha sp.)

 

Conservation status: Near threatened

Disciplines: Animal welfare science, population ecology, community ecology, marine biology, ichthyology

 

Research location: United States


Project summary

Individuals belonging to several elasmobranch populations use the San Francisco Bay Estuary during critical periods of their life history. For example, the broadnose sevengill shark (Notorynchus cepedianus) is an apex predator that visits seasonally to pup. Pups then appear to reside in the estuary for several years before they join the adult population that migrates from Alaska to Baja, California. The San Francisco Bay also serves as a major harbor for container and cruise ships. To allow these large vessels to enter the bay, dredging occurs continuously within San Francisco Bay in the primary channel where many of the adult elasmobranchs also reside, potentially increasing contaminant exposure. Additionally, both commercial and recreational fishing occurs on shark species and their preferred prey, posing direct physiological impacts on elasmobranchs and constraints to their prey availability. Little has been done to evaluate potential unrecognized consequences of these activities on the health and welfare of elasmobranch species and the San Francisco Bay ecosystem. To address this gap in knowledge, this project will measure stress physiology and blood contaminants to evaluate whether the health and welfare of elasmobranchs is threatened within San Francisco Bay, and evaluate dietary needs of elasmobranch species within San Francisco Bay to determine whether individuals are threatened with hunger/starvation due to fishing of their preferred prey. Stable isotope analysis of the sharks and potential prey will also be used to more precisely identify their dietary needs and sensitivities, and ultimately map ecological relationships within the ecosystem.

Why we funded this project

We funded this project primarily because it addresses our interest in understanding the welfare implications of ecological system dynamics. The project also focuses on lifestage-specific ecological differences within the focal species, focusing on juveniles, which are the most numerous and often neglected. Additionally, the lead researcher is a PhD student who has an interest in wild animal welfare and has already demonstrated an aptitude for coordinating scientific research projects. The ability to write a compelling proposal at such an early research stage is very promising for their future career prospects.


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Physiological and behavioral effects of nicarbazin on feral urban pigeons (Columba livia)

Grantee: Jessica X. Wright-Lichter

Institution: Tufts University

Project summary

The oral contraceptive Nicarbazin (NCZ) has been shown to be effective in urban pigeons. However, despite evidence that it may limit birds’ ability to cope with high heat, the side-effects of prolonged exposure to NCZ on pigeons have not been studied. In this project, feral pigeons will be treated with NCZ and exposed to increased ambient temperature to assess their responses. Changes in corticosterone levels will be used as a physiological proxy for welfare. The researchers will also look for behavioral correlations by measuring the onset and frequency of heat-mediating behaviors. The behavioral effects of NCZ treatment alone will also be assessed by quantifying impacts on affiliative and aggressive behaviors, social status, and access to feed at bait sites.

Grantee: Jessica X. Wright-Lichter

 

Institution: Tufts University, United States

Grant amount: $30,000

 

Grant type: Small grants

Focal species: Urban pigeon (Columba livia domestica)

 

Conservation status: Least concern

Disciplines: Physiology, ornithology, animal welfare science

 

Research location: United States


Project summary

Nicarbazin (NCZ) is an oral contraceptive that has been shown to be effective in feral urban pigeons (Columba livia). However, although it is EPA-approved and widely marketed, no published studies examine the impact of prolonged exposure to NCZ on the pigeons beyond the contraceptive effect. This is concerning because evidence from agricultural use of the compound (where it was originally developed as an antiparasitic drug) suggests that NCZ exposure may limit the ability of birds to cope with high heat. 

To study the impact of NCZ treatment on pigeon welfare, especially as a function of heat exposure, feral pigeons will be treated with NCZ for three weeks before being exposed to increased ambient temperature for four hours to assess their physiological and behavioral responses compared to a control group. This project will use changes in levels of the hormone corticosterone as a physiological proxy for welfare in exposed pigeons. The researchers will also look for behavioral correlations by measuring the onset and frequency of heat-mediating behaviors (e.g., panting, wing-spreading). The behavioral (and plausibly welfare-relevant) effects of NCZ treatment alone will also be assessed by quantifying impacts on affiliative and aggressive behaviors, social status, and access to feed at bait sites.

Why we funded this project

We funded this project because we see wildlife fertility controlparticularly applied to urban pigeons — as a highly promising near-term intervention. We are especially interested in indirect welfare effects of fertility control (e.g.,on juvenile welfare and survival), but we have also been concerned by the lack of research on direct welfare impacts. This study will focus on direct welfare impacts while also providing some of the first data on NCZ’s effects on pigeon social dynamics, which are likely to be very important determinants of overall welfare. These data could then be used to guide the application of NCZ contraceptives (e.g., their seasonal timing) to maximize pigeon well-being while successfully controlling urban pigeon populations.


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Integrating individual-level juvenile welfare in dynamic habitats across time and space

Grantee: Tom Luhring

Institution: Wichita State University, Texas State University, and Stephen F. Austin University

Project summary

The project will track four populations of juvenile lesser sirens in Eastern Texas within and across years. Sirens’ health is directly affected by their environment through the impacts of resource availability on body condition and growth rates. Furthermore, sirens show strong size-dependent and seasonal shifts in antagonistic behaviors, which lead to acute injuries. This project will use water-borne corticosterone release rates to investigate changes in stress physiology as a function of changes in the environment experienced by the individual (population density, drought severity index, water temperature, pH, conductivity) across time and space to understand coping capacity. This data will also be used to investigate the welfare impact of an established marking technique compared to a novel machine-learning approach.

Grantee: Tom Luhring

 

Institutions: Wichita State University, Texas State University, and Stephen F. Austin University, United States

Grant amount: $162,604

 

Grant type: Challenge grants

Focal species: Sirens (Siren intermedia)

 

Conservation status: Least concern

Disciplines: Herpetology, physiology, climate science

 

Research location: United States


Project summary

The project will track individual juvenile lesser sirens (Siren intermedia) within and across years for four populations in Eastern Texas. The lack of a terrestrial life-stage and severely limited overland dispersal ability means that hydrologically isolated pools function as closed populations, facilitating recaptures and simplifying demographic estimates. Siren health is directly impacted by the effects of the environment (e.g., drought conditions) through the impacts of resource availability on body condition and growth rates. Furthermore, sirens show strong size-dependent and seasonal shifts in antagonistic behaviors such as biting which lead to acute injuries. 

Aquatic amphibians are especially well-suited for the collection of water-borne stress hormones (corticosterone), which offer the least invasive method of evaluating an integrated measure of corticosterone levels that are passively being released through the skin, gills, feces, and urine. This project will use water-borne corticosterone release rates to investigate changes in stress physiology as a function of changes in the environment experienced by the individual (population density, drought severity index, water temperature, pH, conductivity) across time and space to understand coping capacity. These data will also be used to investigate the welfare impact of an established marking technique compared to a novel approach based on pattern recognition by a machine-learning algorithm. 

Why we funded this project

Juvenile mortality is especially high in amphibians, and amphibian welfare in general is a neglected subject. This project should provide proof of concept for a cost-effective approach for assessing welfare at both an individual and population level. The waterborne measurements have the potential to integrate corticosterone over a longer period of time, increasing its reliability as a welfare indicator. Finally, this project will test a novel, non-invasive approach to mark-recapture studies, which could facilitate much better individual-level welfare research for amphibians and other (especially aquatic) animals in the future.


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Evaluating short- and long-term impacts of injury and illness on wild bird welfare

Grantee: Katie LaBarbera

Institution: San Francisco Bay Bird Observatory

Project summary

Surprisingly little is known about how illness and injury impact the welfare and survival of wild animals, as detecting and assessing injuries and tracking animals to determine their fates is challenging. Yet bird banding stations and wildlife rescues require this information to decide whether birds can be ethically released with long-term impairments. This project will use the San Francisco Bay Bird Observatory (SFBBO)’s long-term bird-banding dataset, which spans over 30 years and 100,000 captures, to investigate these questions in wild birds. Bird banding involves close examination of wild individuals who are frequently recaptured over time. With a high rate of recapture, the SFBBO tracks individuals over years, monitoring their injuries and health, and estimating survival.

Grantee: Katie LaBarbera

 

Institution: San Francisco Bay Bird Observatory, United States

Grant amount: $20,000

 

Grant type: Challenge grants

Focal species: Wild birds

 

Conservation status: Least concern

Disciplines: Wildlife rehabilitation, ornithology

 

Research location: United States


Project summary

There is surprisingly little known about how illness and injury impact the experience of wild animals. Studying such patterns can be limited by the challenges of detecting and assessing injuries and then following up to determine individual fate in wild animals. The San Francisco Bay Bird Observatory’s (SFBBO) long-term bird-banding dataset (>30 years and >100,000 captures) is well-suited to address these questions in wild birds.

Bird banding involves close examination of wild individuals, and individuals are frequently recaptured over time. The SFBBO has a high rate of recapture, which allows them to track individuals over years and to estimate survival, tracking the state of injuries and bird health over multiple years. Understanding how injury impacts individual welfare and survival is of both intellectual and practical value. Bird banding stations vary considerably in their criteria for deciding whether an injured bird should be released or taken to a wildlife rescue. Wildlife rescues must in turn decide whether birds can be ethically released with long-term impairments; for example, many rescues will euthanize rather than release one-legged songbirds. Banding stations and wildlife rescues need real data on wild birds' experiences and prognoses to inform such policies; otherwise, they risk enacting harm.

Why we funded this project

With thousands of wild animal rehabilitation centers in the US alone, this study could provide information that would allow wild animal rehab staff to make data-driven decisions about their bird patients. We think there may be potential to greatly grow interest in the wild animal welfare community via connections with wild animal rehabilitation groups, and this project could provide connections to that community. The project also advances one of our core goals — understanding what wild animals’ lives are like — using an existing and humanely acquired dataset, by providing data on injury rates, severity, and recovery processes. The data could be used to define a metric of “time spent suffering” for injured songbirds.


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Determining the practical and statistical methods necessary for employing field-based metrics of welfare on wild, juvenile, birds

Grantee: Daniel Hanley

Institution: George Mason University

Project summary

Because welfare can vary between individuals and throughout the life of an animal, methods for measuring, assessing, and comparing welfare have been a barrier to our understanding of juvenile welfare. Initial investigations of welfare metrics are needed to estimate age-specific welfare in wild juvenile animals, to determine how they deviate from population-level estimates, and to extend methods and metrics to other systems. This study will examine welfare in free-living prothonotary warblers to establish standardized field and analytical procedures necessary to obtain age-specific animal welfare estimates. Prothonotary warblers are an ideal model system for studying age-specific welfare because they have well-defined life stages, face unique environmental risks, have variable survival, and nest within cavities, affording a degree of standardization and control.

Grantee: Daniel Hanley

 

Institution: George Mason University, United States

Grant amount: $60,000

 

Grant type: Challenge grants

Focal species: Prothonotary warbler (Protonotaria citrea)

 

Conservation status: Least concern

Disciplines: Ornithology, animal behavior, population ecology

 

Research location: United States


Project summary

The juvenile stage is where welfare conditions are likely the most variable and impactful on an individual’s growth and behavior. Unfortunately, methods for measuring, assessing, and comparing welfare have been a barrier to our understanding of juvenile welfare. Like other aspects of animal life history, welfare will vary between individuals and also over the lives of animals in an age-specific fashion. Thus, metrics such as welfare expectancy can inform us of the welfare that an organism is likely to experience, similar to how life expectancy can provide an estimate on how much longer an organism may live. 

This study examines welfare in free-living juvenile songbirds to establish standardized field and analytical procedures necessary to obtain age-specific animal welfare estimates. Prothonotary warblers are an ideal model system for studying age-specific welfare because they have well-defined life stages (i.e., egg, nestling, fledgling, subadult, adult), face unique environmental risks (e.g., drought and flooding), and have variable survival. Furthermore, members of this species nest within cavities, which affords a degree of standardization and control necessary for an initial investigation of welfare metrics. Such initial investigations are crucial to estimate age-specific welfare on wild juvenile animals, to determine how they deviate from population-level welfare estimates, and to extend these methods and metrics to other systems.

Why we funded this project

We funded this project because it sought to explicitly quantify welfare across life stages, using multiple physiological, behavioral, and environmental/demographic indicators. Knowing how (and ideally why) average welfare differs over the course of life in a population could have important implications for interventions to improve their welfare (e.g., fertility control). We were also impressed with this PI because he engages numerous students in their lab and is relatively early in his own career, potentially allowing for pivot to focus more on wild animal welfare. He also demonstrated a good understanding of Wild Animal Initiative’s research on the welfare expectancy framework and sought to put the concepts into practice. That sort of theory-to-practice pipeline would represent a significant step for welfare biology as a research field.


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Thermal imaging to investigate physiological state in altricial nestlings

Grantee: Paul Jerem

Institution: Tufts University

Project summary

Animal stress responses evolved to increase survival, in part by stimulating behaviors that reduce exposure to challenging situations. However, young birds who are entirely reliant on their parents (“altricial”) are incapable of acting to change their circumstances, potentially exposing them to the damaging effects of chronic stress. Such species are known to suppress aspects of their stress physiology during development. However, it remains unclear if other parts of the system remain active and could serve as useful indicators for efforts to improve early life welfare. This project seeks to investigate this possibility in juvenile house sparrows (Passer domesticus) using a novel, non-invasive method for inferring internal state — thermal imaging of body surface temperatures.

Grantee: Paul Jerem

 

Institution: Tufts University, United States

Grant amount: $37,780

 

Grant type: Challenge grants

Focal species: House sparrow (Passer domesticus)

 

Conservation status: Least concern

Disciplines: Physiology, ornithology

 

Research location: United States


Project summary

Animal stress responses evolved to increase survival, in part by stimulating behaviors that reduce exposure to challenging situations. However, young birds who are entirely reliant on their parents (“altricial”) are incapable of acting to change their circumstances, potentially exposing them to the damaging effects of chronic stress. Such species are known to suppress aspects of their stress physiology during development. However, it remains unclear if other parts of the system remain active and could serve as useful indicators for efforts to improve early life welfare. This project seeks to investigate this possibility in juvenile house sparrows (Passer domesticus) using a novel, non-invasive method for inferring internal state — thermal imaging of body surface temperatures.

Why we funded this project

We especially want to understand juvenile welfare, because in most species that is the most experienced life stage, and often the most vulnerable. However, developmental changes can make it difficult to compare welfare between juveniles and adults. This project will calibrate a non-invasive proxy of welfare that can be applied to both adult and altricial juvenile birds, enabling not only better welfare assessments, but more effective lifetime comparisons. A better understanding of the “shape” of lifetime welfare would in turn inform interventions that might disproportionately affect individuals of certain ages.


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The impact of road noise on the welfare of free-living juvenile white-footed mice

Grantee: Michael Sheriff

Institution: University of Massachusetts Dartmouth

Project summary

This project will examine how road noise impacts the ability of juvenile white-footed mice in Massachusetts to respond to the threat of predation. Preliminary work has shown that experimental manipulation of road noise disrupts the foraging responses of (adult) small mammals to predation risk, possibly by masking their ability to perceive predators’ auditory cues. Perception of predation threat will be experimentally manipulated by auditory playback of owl noises at sites near and far from the highway, paired with controls at the same distances from the highway without auditory playback. Anxiety-related behaviors will be recorded in juveniles in an open field trap, and their feces will be studied to assess physiological stress and nutritional status.

Grantee: Michael Sheriff

 

Institution: University of Massachusetts Dartmouth, United States

Grant amount: $60,000

 

Grant type: Challenge grants

Focal species: White-footed mice (Peromyscus leucopus)

 

Conservation status: Least concern

Disciplines: Human-wildlife conflict, animal behavior, population ecology, mammalogy

 

Research location: United States


Project summary

This project will examine the impact of road noise on juvenile welfare in white-footed mice in Massachusetts. The project will focus on how road noise impacts the ability of juveniles to respond appropriately to the threat of predation (the most common cause of juvenile small mammal mortality). Preliminary work has shown that experimental manipulation of road noise (played at 62-65dB, which is equivalent to 100m into the forest from a major thoroughfare to Boston, MA) disrupts the normal foraging responses of (adult) small mammals to predation risk, possibly by masking their ability to perceive auditory cues of predators. Perception of predation threat will be experimentally manipulated by auditory playback of owl noises at sites near and far from the highway, paired with controls at the same distances from the highway but without auditory playback. Anxiety-related behaviors will be recorded in juveniles in an open field trap (which they voluntarily enter for feed), and their feces will be studied to assess physiological stress and nutritional status.

Why we funded this project

Road noise has dramatically increased and is potentially a major anthropogenic threat to wild animal welfare, and one which might be easily ameliorated through policy changes (e.g., improved sound barriers). This project is especially interesting because it focuses on a less obvious effect of road noise, potentially increasing the risk of predation by masking predator cues. This becomes even more interesting in the context of growing literature on the “ecology of fear,” sublethal effects of predators on prey behavior. If road noise makes prey unaware of risks, it could actually reduce their chronic stress despite exposing them to greater risk of death. We are excited for this project to explore those issues, although we are prepared for a complex result. Additionally, we wanted to support this PI because of their strong record of engaging students in their research and influencing their career trajectories.


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Improving the welfare of farmland invertebrates

Grantee: Dr. Ruth Feber

Institution: University of Oxford

Project summary

In Lepidoptera (butterflies and moths), larvae are much more abundant and less mobile than adults. Larvae are therefore particularly vulnerable to negative stimuli, including starvation and disease. This project will use lepidopteran larvae as a model for auditing the welfare impact of agricultural activities on invertebrates. Juvenile stages of Lepidoptera are exposed to agricultural practices that have the potential to affect their welfare. To quantify these impacts, the study will extend the Quality Adjusted Life Year (QALY) methodology to take into account the number of individuals affected by a specified action.

Grantee: Dr. Ruth Feber

 

Institution: Wildlife Conservation Research Unit, Recanati-Kaplan Centre, Department of Zoology, University of Oxford, United Kingdom

Grant amount: $58,448

 

Grant type: Challenge grant

Focal species: Butterflies (Rhopalocera sp.)

 

Conservation status: Near threatened

Disciplines: Entomology, population ecology, physiology

 

Research location: United Kingdom


Project summary

Invertebrates, particularly insects, often have complex life histories. Juveniles (which make up the overwhelming majority of invertebrate numbers) may experience a range of different life quality outcomes. In Lepidoptera (butterflies and moths), for example, adults are the most visible stage, but the much more abundant larvae are less mobile than adults and are particularly vulnerable to negative stimuli including starvation and disease.

This project will use lepidopteran larvae as a model for auditing the welfare impact of agricultural activities on invertebrates. Juvenile stages of Lepidoptera tend to comprise the largest proportion of the total lifespan in temperate regions and, as juveniles, they are exposed to a wide range of agricultural practices that have the potential to affect their welfare. Lepidoptera are also among the better-studied invertebrates, with published data on the ecology, life histories, and survivorship of some species. This knowledge will be used to help inform welfare impact assessments.

The study will adapt the Quality Adjusted Life Year (QALY) methodology to quantify the welfare impacts of agriculture, which has recently been adapted by Teng et al. (2018) to compare the impact of diseases of domestic animals with a Welfare Adjusted Life Year (WALY). This project aims to extend the QALY to take into account the number of individuals affected by a specified action.

Why we funded this project

Farms take up nearly half of the world’s habitable land, but there is a lack of research into how agricultural management practices might impact wild animals, especially invertebrates. In order to improve welfare for invertebrates, we first need to understand how to measure welfare. This project will explore a model to quantify wild insect health and well-being. We were especially attracted to this project because it will repurpose existing data, allowing the research objectives to be accomplished more cheaply and with less animal suffering than might otherwise be required. We were also excited by the PI’s interest in quantifying welfare using a QALY-like framework, which fits perfectly with our utilitarian approach and could lead to actionable policy recommendations.


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Drivers of welfare in American pikas (Ochotona princeps)

Grantee: Johanna Varner

Institution: Colorado Mesa University

Project summary

Pikas (Ochotona princeps) in alpine habitats have recently faced population losses and range retractions. Declines are most prominent in isolated mountain ranges with limited high-elevation habitat, like the La Sals, where pikas are consequently likely to experience lower welfare. This project will assess the welfare of pikas in the La Sals through patterns of survival, population demographics, physiological stress, body condition, ectoparasite load, and hydration. The researchers will also test hypotheses correlating these indicators to habitat quality, and investigate resource competition between pikas and mountain goats.

Grantee: Johanna Varner

 

Institution: Colorado Mesa University, United States

Grant amount: $28,500

 

Grant type: Small grants

Focal Species: American pikas (Ochotona princeps)

 

Conservation status: Least concern

Disciplines: Population ecology, animal behavior, ecological modeling, community ecology

 

Research location: United States


Project summary

Due to recent population losses and range retractions, pikas (Ochotona princeps) are considered a sentinel for climate change in the alpine. However, declines are most prominent in isolated mountain ranges with limited high-elevation habitat, like the La Sals. Surrounded by low-elevation desert in all directions, pikas in the La Sals are likely to experience lower aggregate welfare (i.e., steeper population declines, greater physiological stress, and lower survivorship). This project will assess the welfare of pikas in the La Sals through patterns of survival, population demographics, physiological stress (fecal glucocorticoid metabolites; FGM), body condition, ectoparasite load, and hydration. The researchers will also test hypotheses correlating these welfare indicators to aspects of habitat quality. Finally, they will also investigate resource competition between pikas and mountain goats as another potential factor in pika welfare.

Why we funded this project

This project addresses our research priority of understanding the welfare implications of competition and other conflicts of interest between wild animals. It also focuses on a highly numerous species, the American pika. In that context, we appreciate that this project will model habitat quality on the spatial scale relevant to individual pika when assessing its association with their welfare. Finally, we value that they will use a wide variety of welfare indicators, representing the domains of physiology, behavior, and environment/demography, with each strengthening interpretation of the other.


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