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 continuously update this page as new projects are added.

Does DNA methylation reflect environmental and social adversity?

Grantee: Daniel T. Blumstein

Institution: University of California, Los Angeles

Project summary

This project will investigate the welfare effects of early cumulative adversity in free-living yellow-bellied marmots. It will look for associations between the adversity index and behaviors that indicate general wariness (flight initiation distance, time allocated to vigilance while foraging, the propensity to emit alarm calls while foraging); biomarkers that indicate physiological stress (fecal glucocorticoid levels, neutrophil:lymphocyte ratios); and two measures of aging (telomere length and DNA methylation). By quantifying these behavioral indices of wariness and the suite of biomarkers that culminate in telomeres and epigenetic state in pups throughout their first year and in older animals throughout their lives, the project will determine whether adverse environmental experiences have immediate and lasting effects on welfare.

Grantees: Daniel T. Blumstein, Emily Renkey

 

Institution: University of California, Los Angeles, US

Grant amount: $219,900

 

Grant type: Challenge grants

Focal species: Yellow-bellied marmot (Marmota flaviventer)

 

Conservation status: Least concern

Disciplines: Population ecology, genetics/genomics, ecological modeling, physiology

 

Research locations: United States


Project summary

In humans, early cumulative adversity has demonstrable consequences for health, welfare, and longevity. The welfare consequences of early adversity in wild animals is less well understood, possibly reflecting insufficient measures for assessment. Using a recently validated approach to quantifying cumulative adversity in free-living yellow-bellied marmots (Marmota flaviventer), this project will investigate the welfare effects of early cumulative adversity. It will look for associations between the adversity index and key behaviors that indicate general wariness (flight initiation distance, time allocated to vigilance while foraging, and the propensity to emit alarm calls while foraging), biomarkers that indicate physiological stress (fecal glucocorticoid levels, neutrophil:lymphocyte ratios), and ultimately, two measures of aging (telomere length and DNA methylation). By quantifying these behavioral indices of wariness and the suite of biomarkers that culminate in telomeres and epigenetic state in pups throughout their first year and in older animals throughout their lives, the project will determine whether adverse environmental experiences have immediate and lasting effects on welfare.

Why we funded this project

This project will add welfare to the research portfolio of a long-running study system of a free-living mammal. It will contribute to understanding the validity of biological aging as a welfare indicator by pairing it with other indicators and a comprehensive dataset of the animals’ adverse early-life experiences. 


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Between freedom and security, what makes a feral pigeon happier?

Grantee: Brandon Mak

Institution: Technical University of Munich

Project summary

Feral pigeons in Germany are subject to the Augsburg model of non-lethal population control, which involves attracting them to breed in dovecots where food and nest sites are provided, and their eggs exchanged with dummies. Others are captured and housed in aviaries under similar conditions. This project investigates the welfare implications of Germany’s non-lethal population control strategies. This is done by comparing the stress profiles of feral pigeons living “wild” in the streets, free-living pigeons breeding and feeding in dovecots, and formerly free-living pigeons translocated into aviaries. Fieldwork will be conducted to weigh pigeons (body mass measurement), collect egg and feather samples from their nests (corticosterone concentrations), and observe their behavior (locomotion and aggression).

Grantee: Brandon Mak

 

Institution: Technical University of Munich, Germany

Grant amount: $10,900

 

Grant type: Seed grants

Focal species: Feral pigeon/rock dove (Columba livia domestica)

 

Conservation status: Least concern

Disciplines: Ornithology, animal behavior, physiology, population ecology, wildlife management

 

Research locations: Germany


Project summary

Feral pigeons in Germany are subject to the Augsburg model of non-lethal population control, which involves attracting them to breed in dovecots where food and nest sites are provided, and their eggs exchanged with dummies. Others are captured and housed in aviaries under similar conditions. While aimed at reducing pigeon populations, providing food, and reducing exposure to predators, it’s possible that these measures may incidentally alleviate chronic stress in free-living individuals. This project investigates the welfare implications of Germany’s non-lethal population control strategies. This is done by comparing the stress profiles of feral pigeons living “wild” in the streets, free-living pigeons breeding and feeding in dovecots, and formerly free-living pigeons translocated into aviaries. Fieldwork will be conducted to weigh pigeons (body mass measurement), collect egg and feather samples from their nests (corticosterone concentrations), and observe their behavior (locomotion and aggression).

Why we funded this project

This project will contribute to our understanding of the potential welfare effects of (non-chemical) fertility control in pigeons, a leading candidate for near-term interventions to improve welfare in urban contexts. It will also provide some information on the validity of egg corticosterone as a non-invasive indicator of stress by measuring it alongside other more established methods (feather corticosterone, morphological and behavioural indicators) for a more robust welfare assessment.


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The impact of winter food provisioning on the welfare of wild urban hedgehogs

Grantee: Julia Nowack

Institution: Liverpool John Moores University

Project summary

This project will investigate the welfare impacts of supplemental feeding during winter on European hedgehogs. Hedgehogs and other species have adapted to the lack of natural food during the winter by hibernating. Previous research, however, has suggested that hibernation may be interrupted when anthropogenic food sources are available. This project will use biologgers to collect data on winter activity and hibernation, as well as the animals’ use of specially designed feeding stations equipped with miniature thermal imaging devices, which will be used to non-invasively measure eye temperature as an indicator of physiological stress. Welfare assessment will also be supplemented by data on ectoparasite load, body mass, fecal glucocorticoid concentrations, latency to approach a feeding station, and survival rate for adults and juveniles.

Grantee: Julia Nowack

 

Institution: Liverpool John Moores University, UK

Grant amount: $29,977

 

Grant type: Seed grant

Focal species: Western European hedgehog (Erinaceus europaeus)

 

Conservation status: Least concern

Disciplines: Physiology, population ecology, animal behavior

 

Research locations: United Kingdom


Project summary

This project will investigate the welfare impacts of supplemental feeding during winter on European hedgehogs. Hedgehogs and other species have adapted to the lack of natural food during the winter by hibernating. Previous research, however, has suggested that hibernation may be interrupted when anthropogenic food sources are available. This project will use biologgers to collect data on winter activity and hibernation, as well as the animals’ use of specially designed feeding stations equipped with miniature thermal imaging devices, which will be used to non-invasively measure eye temperature as an indicator of physiological stress. Welfare assessment will also be supplemented by data on ectoparasite load, body mass, fecal glucocorticoid concentrations, latency to approach a feeding station, and survival rate for adults and juveniles.

Why we funded this project

Food provisioning is a potentially important near-term intervention that is already practiced in some contexts, but usually without data on or a primary interest in its wild animal welfare impacts. We appreciate that this project will use a non-invasive approach with multiple physiological and behavioral indicators to evaluate welfare impacts of supplemental feeding on wild European hedgehogs, including juveniles. The results could have implications for the management of other hibernators with access to supplementary feeding, such as dormice, bears, or chipmunks.


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Parasites in the city: How is the impact of parasites on wild animal welfare affected by urbanization?

Grantee: Amanda Trask

Institutions: Institute of Zoology (IOZ), Zoological Society of London (ZSL)

Project summary

This project will evaluate the extent to which parasitism and urbanization impact wild animal welfare, both independently and synergistically, using physiological data on the internal and external parasite loads of deceased gray squirrels, as well as their body condition, immune function, and stress response. The researchers will also collect behavioral welfare indicators from other squirrels using camera traps. This data will be collected along a gradient of urbanization levels and analyzed in the context of the squirrels’ sex and age, to compare population-level welfare across sites varying in degree of urbanization and average parasite metrics and predict how future increases in urbanization might impact wildlife welfare.

Grantee: Amanda Trask

 

Institutions: Institute of Zoology (IOZ), Zoological Society of London (ZSL), United Kingdom

Grant amount: $199,941

 

Grant type: Challenge grant

Focal species: Eastern grey squirrel (Sciurus carolinensis)

 

Conservation status: Least concern

Disciplines: Infectious disease, population ecology

 

Research location: United Kingdom


Project summary

Parasitism is likely a key determiner of wild animal welfare, while rapidly increasing urbanisation presents novel challenges and welfare threats, yet the extent that parasitism and urbanisation impact wild animal welfare, both independently and synergistically, remains a major knowledge gap. An individual’s parasite infection status, and the associated welfare impacts, is dependent on parasite exposure risk and susceptibility. Urbanisation may alter exposure risk, for example through changes to population density and resource distribution, and susceptibility, for example through altered host immune function from exposure to urban stressors. Urbanisation may also compound impacts of parasitism, if chronically stressed individuals are less able to cope with further welfare challenges, resulting in synergistic negative welfare effects. Furthermore, welfare impacts of parasites and urbanisation may vary across sex- and life-stage classes, for example due to hormonal influences and changes in immune function with chronic exposure to urban pollution.

We will address this knowledge gap, using grey squirrels from UK sites across an urbanisation gradient. We will use detailed individual-level data on parasite loads and physical and physiological welfare indicators, collected from dead squirrels killed as part of non-native species management, to determine welfare impacts across sites varying in degree of urbanisation, and across sexes and life stages. Then, we will use behavioural indicators from camera trap data to determine associations between population-level welfare, parasite prevalence and degree of urbanisation. Finally, we will estimate ‘welfare expectancy’ of individuals from different populations using sex- and life-stage specific welfare scores. We will expand on current theory and develop a modelling tool to incorporate demographic and environmental stochasticity in welfare states into population models, to aid prediction of how increasing urbanisation may impact wildlife welfare in the future. This study will provide vital knowledge to aid understanding and prediction of the experience and future responses of wild animals to parasites in the context of increasing urbanization, facilitating design and integration of welfare-friendly urban environments for wildlife.

Why we funded this project

Gray squirrels are abundant, they live in close proximity to humans, and they could benefit from welfare interventions such as wildlife fertility control in the near future. Gray squirrels are not native in the UK, where this study will be conducted, and the welfare of non-native species is generally understudied or undervalued compared with native species of conservation interest. This project provides a special opportunity to take advantage of data from squirrels who are being killed as part of a government program, and to access valuable metrics that are hard to measure non-invasively, thereby creating from their deaths an opportunity to gain valuable insights that could ultimately help other squirrels. In addition to their good range of welfare indicators, we appreciate that this project will consider sex- and age-specific variation in welfare impacts and attempt to estimate “welfare expectancy” at a population level.


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Predicting density dependence of welfare of wild animal populations based on resource access linked to habitat availability and usage

Grantee: Ross MacLeod

Institution: Liverpool John Moores University

Project summary

This project seeks to adapt a habitat and population model to incorporate welfare and apply it to the study of an abundant and widespread avian species, the house sparrow (Passer domesticus). The researchers propose to utilize a variety of welfare metrics to validate model assumptions, which will then allow them to test assumptions related to the welfare implications of density-dependent population dynamics.

Grantee: Ross MacLeod

 

Institutions: Liverpool John Moores University, United Kingdom

Grant amount: $159,744

 

Grant type: Challenge grants

Focal species: House sparrow (Passer domesticus)

 

Conservation status: Least concern

Disciplines: Ecological modeling, population ecology, animal welfare science

 

Research location: United Kingdom


Project summary

This project seeks to adapt a habitat and population model to incorporate welfare and apply it to the study of an abundant and widespread avian species, the house sparrow (Passer domesticus). The researchers propose to utilize a variety of welfare metrics to validate model assumptions, which will then allow them to test assumptions related to the welfare implications of density-dependent population dynamics.

Why we funded this project

We funded this project because it addresses our proposal request very closely and proposes to investigate a key wild animal welfare question using a modeling framework. They are also planning to address their question using an abundant avian species. The project has high potential to inform future work focused on modeling total welfare in a population (i.e., combining both individual welfare and population size), and to create a model that can be replicated in other systems.

Find Ross’ other project, studying wild birds, here.


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Thermal imaging as a non-invasive welfare assessment tool for tracking the impact of environmental stressors across wild animal populations

Grantee: Ross MacLeod

Institution: Liverpool John Moores University

Project summary

This project aims to test and validate a standardized multi-species approach to monitoring physiological stress in wild birds by using thermal imaging cameras to measure the animals’ body surface temperature, which could enable tracking of chronic stress in wild populations facing environmental stressors. Building on pilot work, the project will focus on wild bird populations to develop a standardized thermal imaging methodology capable of monitoring surface temperature of a wide range of wild animals. The methodology will be validated using bird communities visiting 54 standardized feeding and drinking stations spread across the UK, to quantify how changes in surface temperature are linked to starvation risk, predation risk, and human disturbance.

Grantee: Ross MacLeod

 

Institution: Liverpool John Moores University, United Kingdom

Grant amount: $29,810

 

Grant type: Small grants

Focal species: Wild birds

 

Conservation status: Least concern

Disciplines: Physiology, animal welfare science, population ecology, ornithology

 

Research location: United Kingdom


Project summary

This project aims to test and validate a standardized multi-species approach to monitoring physiological stress in wild birds by using thermal imaging cameras to measure the animals’ body surface temperature, which could enable tracking of chronic stress in wild populations facing different levels of environmental stressors. Building on pilot work, the project will focus on wild bird populations to develop a standardized thermal imaging methodology capable of monitoring surface temperature of a wide range of free-living wild animals. The methodology will be validated using bird communities visiting 54 standardized feeding and drinking stations spread across three urban-rural gradients in the UK, to quantify how changes in surface temperature are linked to starvation risk, predation risk, and human disturbance.

Why we funded this project

This approach to measuring wild animals’ physiological stress levels through thermal imaging analysis has great potential for scalable monitoring of large numbers of individuals and is usable for cross-species comparisons. We appreciated that this team was interested in addressing non-anthropogenic causes of suffering in starvation and predation, and in extending their method to other species that tend to be neglected (e.g., wild rodents). This work also relates to the project by Paul Jerem that we previously funded, creating a longer-term relationship between Wild Animal Initiative and project participants.

Find Ross’ other project, studying house sparrows, here.


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Novel epigenetic approaches to measure wild animal welfare and stress

Grantee: Dave Daversa

Institution: University of California, Los Angeles

Project summary

A challenge in wild animal welfare science is developing composite assays that consider the full breadth of factors collectively shaping subjective experiences. This project will demonstrate proof of concept for a DNA methylation (DNAm)-based model of wild animal welfare, characterizing DNAm in western toads. It will test the influence of infection with Batrachochytrium dendrobatidis (Bd) on DNAm and its covariance with demographic factors such as age, sex, and body condition. Captive-reared toads will be sampled to characterize DNAm under controlled conditions, providing a standardized profile of DNAm rates over toad life stages. The researchers will then sample toads from wild populations experiencing starkly different levels of Bd infection, with accelerated biological aging signaling elevated stress and impaired welfare.

Grantee: Dave Daversa

 

Institution: University of California, Los Angeles, United States

Grant amount: $30,000

 

Grant type: Small grants

Focal species: Western toad (Anaxyrus boreas)

 

Conservation status: Least concern

Disciplines: Physiology, herpetology, animal welfare science, population ecology, genetics/genomics

 

Research location: United States


Project summary

A major challenge in wild animal welfare science lies in developing composite assays that consider the full breadth of factors collectively shaping the subjective experiences of animals. This project will apply epigenetic tools to develop minimally invasive and aggregate measures of wild animal welfare. Specifically, the researchers aim to demonstrate proof of concept for a DNA methylation (DNAm)-based model of wild animal welfare. DNAm joins other measurements such as telomere attrition as a biomarker of biological aging, because DNA becomes methylated as a function of both chronological age (time) and accumulated stress. This project will characterize DNAm in western toads (Anaxyrus boreas) from wild populations in southern California, testing the influence of infection with the fungal pathogen Batrachochytrium dendrobatidis (Bd) on DNAm, and its covariance with demographic factors such as age and sex, as well as with body condition, a common metric of physical health in wild animals. Captive-reared toads representing all life stages will be sampled to characterize DNAm under controlled conditions, providing a standardized profile of DNAm rates over toad life stages. The researchers will then sample toads from wild populations experiencing starkly different levels of environmental stress (Bd infection), with accelerated biological aging signaling elevated stress and impaired welfare. 

Project objectives

Objective 1: Determine the relationship between DNAm and basic risk factors linked to welfare: chronological age, sex, weight, and infection status.

Objective 2: Determine the effect of environmental stressors on epigenetic age.

Why we funded this project

We funded this project because we are interested in the potential for biological aging biomarkers to be used as very long-term, integrative metrics of animals’ lifetime welfare. DNAm is one such potential biomarker that has received relatively little attention in a welfare context. However, DNAm may potentially be easier to measure than more commonly discussed measures of biological age — or at least, be more familiar for mainstream ecological genetics researchers — because DNA methylation is already of interest for other reasons in biological science. We were particularly enthusiastic about this project because it focuses on a species belonging to a large species complex of amphibians, including both common and threatened species, offering broad transferability and potential impact. Amphibians are also relatively neglected in terms of welfare research. We appreciate that this project aims to establish baseline age-specific differences in biological aging rate in order to then assess age-specific differences attributable to different exposures (i.e., age-specific differences in welfare), which connects to the concept of “welfare expectancy” developed by Wild Animal Initiative researcher Luke Hecht.

Find Dave’s other project, studying western fence lizards, here.


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Developing an approach for assessing the welfare of wild birds through the use of bioacoustics analysis

Grantee: Luiza Figueiredo Passos

Institution: Liverpool John Moores University

Project summary

Vocalization analysis has been used to monitor the welfare of farmed animals and animals in laboratories, but lacks validation for wild animals. This project will use passive acoustic monitoring to record wild bird calls throughout the UK. It will identify vocal patterns and chorus characteristics related to various environmental risks that can be used to infer the welfare status of local bird populations across multiple environments. The focal species will be house sparrows, who have undergone a massive population decline and may be facing welfare challenges. Validating acoustic monitoring as a tool for non-invasive welfare monitoring in the wild for the first time will provide a method that could be used to monitor the welfare of wild birds globally.

Grantees: Luiza Figueiredo Passos

 

Institution: Liverpool John Moores University, United Kingdom

Grant amount: $29,060

 

Grant type: Small grants

Focal species: House sparrow (Passer domesticus)

 

Conservation status: Least concern

Disciplines: Animal welfare science, ornithology, population ecology, animal behavior

 

Research location: United Kingdom


Project summary

Vocalisations carry emotional, physiological, and individual information, suggesting that they can serve as potentially useful non-invasive indicators for inferring wild animal welfare. Communication of affective states via vocalization is due to changes in emotions leading to deviations in the nervous system, causing physiological changes within the vocal production systems. Modern sound analysis techniques have provided tools to discriminate, analyze, and classify specific vocalizations, permitting them to be used for monitoring welfare of different farm and laboratory animals. However, this approach lacks validation for wild individuals. This proposal aims to use passive acoustic monitoring to record wild bird calls at different locations in urban, rural, and conservation-managed areas in the UK. This project will follow a comparative approach to identify vocal patterns and chorus characteristics related to different environmental risks (starvation risk, predation risk, human disturbance, etc.) so that these vocal signatures can be used to infer the welfare status of local bird populations across different environments. The focal species will be one of Europe’s most common species, the house sparrow (Passer domesticus). House sparrows provide an ideal first validation as the species has undergone a massive population decline in many parts of its range over the past 50 years, suggesting that its local populations are facing significant welfare challenges. Validating acoustic monitoring as a tool for non-invasive welfare monitoring in the wild for the first time will provide the basis for future application to birds in general, providing a method that could be used to monitor welfare and changes in welfare of wild birds at any location globally.

Project objectives

Objective 1: Evaluate how changes in sparrow welfare driven by changes in risk of starvation can be indicated by changes in the vocal signatures of local populations.

Objective 2: Understand relationship between predation risk driven changes in sparrows’ welfare and the vocal signatures of local populations.

Objective 3: Impact of human disturbance on sparrows’ welfare.

Why we funded this project

This project is very similar to projects by Claudia Wascher and Sam Reynolds. As stated, bioacoustic monitoring is an extremely promising approach because it is minimally invasive and inexpensive. However, it still needs to be validated, and given its potential future value, we felt that it would be best for it to be validated by at least two independent studies. This project was especially appealing because it focuses on such a ubiquitous species (the house sparrow) and is led by a researcher with a strong record in bioacoustics and an understanding of animal welfare science.

Find Luiza’s other project, studying wild newts, here.


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Assessing animal welfare from bioacoustic monitoring in red-billed choughs (Pyrrhocorax pyrrhocorax)

Grantee: Claudia Wascher

Institution: Anglia Ruskin University

Project summary

Bioacoustic monitoring is a well established method of measuring welfare in captive animals, but has not been used in wild animals. This project will assess and validate the use of passive bioacoustic monitoring to assess welfare in a reintroduced population of red-billed choughs. The researchers will conduct a combination of automatic acoustic monitoring using “passive” acoustic recorders and direct observations (recording behavior via videos and monitoring activity during the night with night vision scopes) and “active” acoustic monitoring, using a high-quality directional microphone and parabolic dish. Their approach will allow them to record vocalizations and behavior during natural events expected to influence welfare, such as anthropogenic noise, predation events, intraspecific aggression, resting, or affiliative behaviors.

Grantees: Claudia Wascher

 

Institution: Anglia Ruskin University, United Kingdom

Grant amount: $18,679

 

Grant type: Small grants

Focal species: Red-billed chough (Pyrrhocorax pyrrhocorax)

 

Conservation status: Least concern

Disciplines: Animal welfare science, ornithology, population ecology, animal behavior

 

Research location: United Kingdom


Project summary

In principle, animal vocalizations can be used to identify both positive and negative experiences. Specific vocal characteristics (e.g., mean fundamental frequency) and the rate of occurrence of certain call types can help to quantify animal welfare and are well-established in farm animals such as pigs, horses, cows, and chickens. But although bioacoustic monitoring is a well-established method in captive animals, the method has not been used in wild animals to assess positive and negative emotional states. This project will assess and validate the use of passive bioacoustic monitoring to assess welfare in a reintroduced population of red-billed choughs. The researchers will conduct a combination of automatic acoustic monitoring using “passive” acoustic recorders and direct observations (recording behavior via videos, monitoring activity during the night with night vision scopes, and “active” acoustic monitoring using high-quality directional microphone and parabolic dish). Their approach will allow them to record vocalizations and behavior during natural events expected to influence welfare, such as anthropogenic noise, predation events, intraspecific aggression, resting, or affiliative behaviors. 

Why we funded this project

This project will help to validate the use of passive acoustic monitoring to assess welfare states from variation in vocal behavior, which presents a non-invasive, cost-effective tool to assess the impact of positive and negative events on animal welfare. In theory, acoustic monitoring could be used to assess the welfare of a large number of individuals at a time, making it a particularly promising method to investigate.


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Impacts of land-use on social networks in mixed-species bird flocks, with implications for the short-term and long-term welfare of Himalayan birds

Grantee: Akshay Bharadwaj

Institution: Indian Institute of Science

Project summary

This project will use mist-netting and bird-banding, followed by standardized observations of behavior and body condition, to examine the relationship between individual-level social behavior, bird health, and survival within mixed-species bird flocks (MSFs) in Eaglenest Wildlife Sanctuary, Arunachal Pradesh, India. Feather corticosterone level, ectoparasite load, and other morphological parameters will be used to measure the health of each banded individual. The researcher will also take advantage of a long-term bird-banding dataset, spanning 12 years, to examine the linkages between inter-individual differences in social behavior and the survival of each individual, comparing survival rates and reproductive success in primary and logged forest.

Grantee: Akshay Bharadwaj

 

Institutions: Indian Institute of Science, India

Grant amount: $15,000

 

Grant type: Small grants

Focal species: Multi-species birds

 

Conservation status: Least concern

Disciplines: Ornithology, physiology, community ecology, infections disease, population ecology

 

Research location: India


Project summary

This project will use mist-netting and bird-banding, followed by standardized observations of behavior and body condition, to examine the relationship between individual-level social behavior, bird health, and survival within mixed-species bird flocks (MSFs) in Eaglenest Wildlife Sanctuary, Arunachal Pradesh, India. Feather corticosterone level, ectoparasite load, and other morphological parameters will be used to measure the health of each banded individual. The researcher will also take advantage of a long-term bird-banding dataset, spanning 12 years, to examine the linkages between inter-individual differences in social behavior and the survival of each individual, comparing survival rates and reproductive success in primary and logged forest. 

Why we funded this project

Understanding the importance of multi-species flocking behavior in birds is relevant to our research priority of understanding indirect welfare effects in ecological systems, which this project approaches in a cost-effective way. We were especially impressed by the quality of the proposal for this project, particularly as it is led by a beginning graduate student in India. Funding this project serves to increase the geographic diversity of our grantee community and therefore of the nascent field of wild animal welfare research.


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