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.

Integrating behavioral competency and post-release support for reintroduced wildlife: a shift in paradigm for rehabilitation and beyond

Grantee: Karli Rice Chudeau

Institutions: The Marine Mammal Center, University of California, Davis

Project summary

This project investigates post-release support and monitoring to improve outcomes for rehabilitated juvenile pinnipeds. Post-release support will include familiar cognitive enrichment to help released animals adjust gradually and buffer their affective state. Post-release monitoring will consider metrics such as behavioral diversity, energy expenditure, and body condition, and the animals’ specific behavioral profiles will be compared with those recorded from healthy, wild individuals. These metrics will be used to evaluate the effectiveness of post-release enrichment as an intervention for improving welfare outcomes. Cognitive bias tests for affective state carried out during rehabilitation and prior to release will also be considered as potential predictors of post-release welfare.

Grantee: Karli Rice Chudeau

 

Institutions: The Marine Mammal Center, University of California, Davis, United States

Grant amount: $30,000

 

Grant type: Seed grants

Focal species: Northern elephant seal (Mirounga angustirostris), eastern Pacific harbor seal (Phoca vitulina richardii)

 

Conservation status: Least concern

Disciplines: Wildlife rehabilitation, animal behavior, animal welfare science

 

Research location: United States


Project summary

In many cases, the process of releasing a rehabilitated or translocated animal can be traumatic and removes the animal’s agency, potentially weakening their ability to thrive in the wild. This project investigates post-release support and monitoring to improve outcomes for rehabilitated juvenile pinnipeds. Post-release support will include familiar cognitive enrichment to help released animals adjust gradually and buffer their affective state. Post-release monitoring will consider metrics such as behavioral diversity, energy expenditure (distance traveled), and body condition, and the animals’ specific behavioral profiles (e.g. foraging behavior) will be compared with those recorded from healthy, wild individuals. These metrics will be used to evaluate the effectiveness of post-release enrichment as an intervention for improving welfare outcomes. Cognitive bias tests for affective state carried out during rehabilitation and prior to release will also be considered as potential predictors of post-release welfare.

Why we funded this project

We envision a world in which people take responsibility for improving wild animals’ lives and have the knowledge they need to do so effectively. Rehabilitation is a part of that. However, there has been relatively little research on post-release outcomes for rehabilitated animals. Understanding those outcomes and identifying strategies to improve them could have significant welfare implications, especially for the treatment of juvenile animals, whose life trajectories may be powerfully affected by the rehabilitation and release process. We appreciate that this project combines post-release monitoring with both a specific intervention and pre-release tests of affective state that would not be possible in a wild context.


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Social connections and their welfare implications in the wild

Grantee: Alex Thornton

Institutions: University of Exeter, University of Bristol

Project summary

This project seeks to understand the relationship between welfare and social interactions in wild bird populations. Using historical data, the researchers will also seek insight into how welfare varies among individuals in relation to the social system, early-life experiences, and interactions among individuals. They will also investigate whether social systems might play a role in helping to mitigate some of the negative anthropogenic impacts on welfare.

Grantee: Alex Thornton

 

Institutions: University of Exeter, University of Bristol, United Kingdom

Grant amount: $157,962

 

Grant type: Challenge grants

Focal species: Jackdaws (Coloeus sp.)

 

Conservation status: Least concern

Disciplines: Animal behavior, ornithology, animal welfare science

 

Research location: United Kingdom


Project summary

This project seeks to understand the relationship between welfare and social interactions in wild bird populations. Using historical data, the researchers will also seek insight into how welfare varies among individuals in relation to the social system, early-life experiences, and interactions among individuals. They will also investigate whether social systems might play a role in helping to mitigate some of the negative anthropogenic impacts on welfare.

Why we funded this project

The study findings will help in understanding how wild animals cope with increasing temperatures and the impact of thermal stress on their welfare and health. Notably, previous work has suggested that T3 measurements in ungulates are especially sensitive to thermal stress, and so comparing T3 with other indicators based on different physiological pathways, such as glucocorticoids, could help researchers to diagnose the relative significance of different environmental stressors an animal is facing. The project’s behavioral metrics are also crucial for realizing that potential. A secondary reason for our interest in this project is that it has near-term policy implications, potentially highlighting the value of preserving or promoting specific landscape features for the ecosystem service they offer, in the form of shade, to wild ungulates.


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Paying to play? Social time allocation, emotional contagion, and social attention as assays of welfare in wild animals

Grantee: Patrick Tkaczynski

Institution: Liverpool John Moores University

Project summary

Play and grooming have a number of facets that could be potential indicators of positive welfare in group-living vertebrates, but analysis of such behaviors are yet to be put into a formal framework for monitoring wild animal populations. This pilot project will focus on six different wild Barbary macaque groups of variable sizes and facing varying degrees of human exposure, which is typically associated with poor health outcomes in wild animals. It will use play and grooming behavior to quantify social time allocation, emotional contagion/synchronization, and social attention as potential assays of positive welfare. Once validated, these assays can be applied to examine how positive welfare varies across different gradients, such as climate change or variation in predator pressure.

Grantee: Patrick Tkaczynski

 

Institution: Liverpool John Moores University, United Kingdom

Grant amount: $29,323

 

Grant type: Small grants

Focal species: Barbary macaque (Macaca sylvanus)

 

Conservation status: Endangered

Disciplines: Animal welfare science, animal behavior

 

Research location: Morocco, United Kingdom


Project summary

For group-living vertebrates, play and grooming have a number of facets that could be potential indicators of positive welfare, but analysis of such behaviors are yet to be put into a formal framework for monitoring wild animal populations. The Middle Atlas region of Morocco hosts one of the largest populations of wild Barbary macaques, an endangered primate species. These animals face a number of environmental challenges that may disrupt positive social experiences, including anthropogenic disturbance, threats from natural and introduced predators, and an extreme climate. Focusing on six different macaque groups with variable group sizes and facing varying degrees of human exposure, this pilot project will use play and grooming behavior to quantify social time allocation, emotional contagion/synchronization, and social attention as potential assays of positive welfare. As human exposure is typically associated with poor health outcomes in wild animals, this pilot project will focus on this specific environmental challenge to help validate our assays, which can later be applied to examine how positive welfare varies across different gradients, such as climate change or variation in predator pressure. 

Why we funded this project

Most animal welfare research is focused on minimizing suffering, but it’s also vital to maximize positive experiences. The links between positive welfare and evolutionary fitness are also less obvious than for negative welfare, where factors like poor health seem to be associated with both poor welfare and low evolutionary fitness. Because more research focuses on fitness than welfare, the unclear relationship between fitness and positive welfare has contributed to the neglectedness of indicators of positive welfare. This project was one of the few we received that addressed the theme of positive welfare, and we hope that this pilot in primates will provide information to support similar studies in other social species.


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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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A bird’s eye view to the five domains of welfare: a quantitative framework and proof-of-concept evaluation in a cetacean, Orcinus orca

Grantees: Saana Isojunno, Eve Jourdain

Institutions: University of St. Andrews, Norwegian Orca Survey

Project summary

This project will carry out body condition and welfare assessments for inshore-foraging killer whales in the northeast Atlantic. The researchers will use the Five Domains model to categorize likely factors influencing killer whale welfare and quantify some of those factors using aerial photography from drones. They will gather data on foraging time and feeding rates, group composition and surface behavior, body shape as a proxy for body condition and blubber reserves, and reproductive success. The researchers will then develop a proof-of-concept statistical model to infer latent motivational states beneath the observable data. The project will also utilize data from known cases of poor welfare to help define their welfare scale.

Grantees: Saana Isojunno and Eve Jourdain

 

Institutions: University of St. Andrews, Scotland, and Norwegian Orca Survey, Norway

Grant amount: $28,707

 

Grant type: Small grants

Focal species: Orca (Orcinus orca)

 

Conservation status: Data deficient

Disciplines: Animal welfare science, bio/eco-informatics, physiology, animal behavior, marine biology

 

Research location: Scotland, Norway


Project summary

This project will carry out body condition and welfare assessment for inshore-foraging killer whales (Orcinus orca) in the northeast Atlantic as an extension to ongoing research in Scotland and Norway. The researchers will use the Five Domains model to categorize likely factors influencing killer whale welfare, and then quantify some of those factors using aerial photography from drones. Specifically, the researchers intend to gather data on foraging time and feeding rates (nutrition), group composition and surface behavior (behavior), and body shape as a proxy for body condition and blubber reserves (health). Data will also be collected on reproductive success (e.g., calf loss). The researchers will then develop a proof-of-concept statistical model to infer latent motivational states (i.e., “true” welfare as a psychological state) beneath the observable data. The project will also utilize data from known cases of poor welfare (where individual social and nutritional needs are not met) to help define their welfare scale.

Why we funded this project

We funded this project because we see hidden state models as promising statistical tools for representing the relationship between disparate data types and welfare, and would like to see this project provide a proof of the concept. This project will also implement a cost-effective and non-invasive approach method based on photogrammetry to assess cetacean health and behavior. Finally, we wanted to fund this project because it spans multiple universities and a citizen science community, creating significant networking opportunities to promote welfare biology.


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Mapping welfare for conservation management

Grantee: Lisa Olivier

Institution: Game Rangers International

Project summary

This project will monitor behavioral and physiological parameters in African elephants over one year to map their perception of their environment. The aim is to test this approach’s potential as a leading indicator that is easy to measure, responds quickly to ecosystem changes, and highlights potential detrimental impacts. These measures will be aggregated to define a scale representing the animals’ perceptions of threat that can be estimated for a given time and place. The researchers will then project this data onto maps to identify areas of high and low welfare. The project will help conservation practitioners create management plans to protect wild animals by providing a head start in assessing changes in their surroundings.

Grantee: Lisa Olivier

 

Institutions: Game Rangers International, Zambia

Grant amount: $29,791

 

Grant type: Small grants

Focal species: African elephant (Loxodonta)

 

Conservation status: Endangered

Disciplines: Animal behavior, animal welfare science, wildlife management

 

Research location: Zambia


Project summary

To effectively steward wild animal welfare, leading indicators are needed that predict the future rather than summarizing the past.  These must be easy to measure, respond quickly to ecosystem changes and highlight potential detrimental impacts that need addressing, including anthropogenic activities. This project aims to test the approach of mapping wild animals’ perception of their environment by monitoring an array of behavioral and physiological parameters in African elephants over a year as they navigate the landscape. These measures will be aggregated to define a scale representing the animals’ perceptions of threat that can be estimated for a given time and place. The researchers will then project these data onto maps to identify areas of high and low welfare, where animals may be feeling alternatively secure or fearful. The researchers intend for the project to help conservation practitioners create better management plans to protect wild animal populations by providing a head start in assessing changes in their surroundings.

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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Every breath you take, I‘ll be watching you: Automated measurement of breath rate from mobile phone videos as a severity assessment parameter in wild great tits

Grantee: Caroline Deimel

Institution: Max Planck Institute

Project summary

This project aims to provide a validated reference tool for standardizing breath rate (BR) measurements following capture and physical sampling protocols in wild birds. The researchers will use computer vision capabilities and other recently developed software improvements to estimate BR metrics from mobile phone videos, enabling the collection of objective, reproducible, and comparable data, and providing institutions tasked to oversee animal welfare with objective and feasible monitoring requirements. The project will also evaluate BR as a welfare indicator in great tits by analyzing an existing five-year dataset to test whether BR corresponds to simultaneous glucocorticoid measurements from free-living great tits. This data will provide reliable baselines and ranges for glucocorticoids and BR, and benchmarks for video recording lengths.

Grantee: Caroline Deimel

 

Institutions: Max Planck Institute, Germany

Grant amount: $19,200

 

Grant type: Small grants

Focal species: Great tits (Parus major)

 

Conservation status: Least concern

Disciplines: Animal behavior, animal welfare science, physiology

 

Research location: Germany


Project summary

Breath rate (BR) is increasingly used as a non-invasive proxy of stress that is fast, cheap, and field-friendly. However, BR has not been evaluated in a bird welfare context, and it is unclear how it relates to established physiological proxies of stress, like glucocorticoid measurements in blood. Also, the currently used protocols to measure BR lack standardization, scalability, and validation. This project aims to provide a validated, non-invasive reference tool for standardizing BR measurements following capture and physical sampling protocols in wild birds by implementing computer vision capabilities and other improvements in software the researchers have recently developed to estimate BR metrics from mobile phone videos. This will allow the research community to gather objective, reproducible, and comparable data, and provide institutions tasked to oversee animal welfare with objective and feasible monitoring requirements. The project will also evaluate BR as a welfare indicator in great tits (Parus major), a songbird extensively used in wild animal research across Europe, by analyzing an existing, five-year dataset to test whether BR corresponds to simultaneous glucocorticoid measurements from free-living great tits. These data will provide reliable baselines and ranges for glucocorticoids and BR, and benchmarks for video recording lengths. 

Why we funded this project

We are generally interested in developing non-invasive ways of measuring indicators of wild animal welfare. One of the key advantages to non-invasive measurement, besides the obvious of not causing unnecessary fear or pain to animals, is that stress induced by the measurement process can obscure the animal’s baseline stress levels if the method is too invasive or not carried out properly. We also tentatively believe that instilling a norm of minimizing animal harm within welfare biology research will increase the likelihood that researchers act as scientist-advocates for implementation of wild animal welfare interventions. This project’s focus on making breath rate easy and inexpensive to measure in a consistent way also fits well with our desire for a greater volume and accessibility of welfare data collection.


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City living – assessing the welfare costs of urban living in spotted hyaena (Crocuta crocuta)

Grantee: Emma Stone

Institutions: Carnivore Research Malawi, University of the West of England

Project summary

Networks of green spaces and river corridors support a population of urban spotted hyenas in Lilongwe City, Malawi. However, human-wildlife conflict (HWC) is frequent. Using non-invasive techniques, this project will measure and compare the fecal and hair cortisol levels, mean body condition, tooth ware, hematological parameters, and disease in urban and rural populations of spotted hyenas. Behavioral experiments (exposure to novel objects and threatening scents) will also be used to assess individual boldness in urban and rural populations as a risk factor, which may increase propensity for HWC and therefore mortality and stress. Results will be used to inform policies to reduce HWC and implement welfare considerations as part of the population’s conservation management plan.

Grantee: Emma Stone

 

Institutions: Carnivore Research Malawi and University of the West of England, United Kingdom

Grant amount: $30,000

 

Grant type: Small grants

Focal species: Spotted hyaena (Crocuta crocuta)

 

Conservation status: Least concern

Disciplines: Human-wildlife conflict, physiology, animal welfare science, animal behavior, infections disease

 

Research location: Malawi


Project summary

Lilongwe City (the capital of Malawi, central Africa) has a good network of green spaces and river corridors, supporting a population of urban spotted hyena (Crocuta crocuta). However, human-wildlife conflict (HWC) is frequent, often resulting in persecution of these animals. Using non-invasive techniques, this project will measure and compare the fecal and hair cortisol levels (as a proxy of stress), mean body condition, tooth ware, hematological parameters, and disease (toxoplasmosis and rabies) in spotted hyena between urban (Lilongwe district) and rural (Kasungu National Park) populations. Behavioral experiments (exposure to novel objects and threatening scents) will also be used to assess individual boldness in urban and rural populations as a risk factor, which may increase propensity for HWC and therefore mortality and stress. Results will be used to inform policies to reduce HWC and implement welfare considerations as part of the population’s conservation management plan.

Why we funded this project

Interactions between people and wildlife are increasing as urban areas expand. This is of particular concern with carnivores, as even small species can be perceived as threatening. Despite the fact that the developing world has the highest rate of urbanization, urban carnivore studies have generally focused on European and Asian cities, with research on African cities being relatively neglected. Although HWC is a traditional issue in conservation, we have encountered few projects that effectively address the more neglected question of how HWC interacts with non-anthropogenic welfare impacts. By considering a variety of welfare indicators/factors, including disease transmission and social dynamics, this project will help us understand that interaction, producing results that can be translated into more abundant urban species. Additionally, by funding at least some projects on endangered species, we increase the likelihood of our welfare-focused message reaching conservation practitioners, potentially leading to a greater emphasis on and interest in welfare within that community.


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WellSeal: Non-invasive quantification of welfare in lactating harbour seals

Grantee: Jess Harvey-Carroll

Institution: University of Gothenburg

Project summary

Limited availability of harbor seal birthing sites leads to intraspecific competition, forcing mothers to choose when and for how long to forage, and resulting in trade-offs between maternal and offspring welfare. These trade-offs may be exacerbated by fishing, as harbor seals swim large distances to obtain food. This project will use camera trap footage and 3D modeling to assess the influence of maternal welfare on time spent away from pups, and pup welfare during periods of maternal absence. Welfare will be assessed based on body condition, an index of the optimality of haul-out position, and a record of positive and negative interactions. The assessment will be aggregated at colony level to compare welfare between colonies with varying access to food.

Grantee: Jess Harvey-Carroll

 

Institution: University of Gothenburg, Sweden

Grant amount: $30,000

 

Grant type: Small grants

Focal species: Harbour seal (Phoca vitulina)

 

Conservation status: Least concern

Disciplines: Population ecology, animal behavior, animal welfare science

 

Research location: Sweden


Project summary

Harbor seals require locations protected from adverse weather and other disturbances during the birthing season, when vulnerable pups are left on land while mothers forage. The limited availability of such sites leads to high intraspecific competition, in which larger seals are likely to dominate more desirable locations due both to their physical strength and their lower need to forage. Mothers must choose when and for how long to forage, leading to trade-offs between maternal and offspring welfare which may lay the foundation for the welfare of younger generations. Currently, little information is available on time spent foraging during the lactation period. As harbor seals are facing unprecedented stress from overfishing, seals are swimming increasingly large distances to obtain food. It is not known how this change affects the trade-off between maternal foraging and offspring care, and the resulting welfare of the pups. This project will use camera trap footage and state-of-the-art 3D modeling to non-invasively assess the influence of maternal welfare on the time spent away from the pups. Pup welfare during periods of maternal absence will also be assessed. Welfare will be assessed in terms of the Five Domains framework, based on individual body condition (health), an index of the optimality of their haul-out position (environment), and a record of positive and negative interactions (behavior). Such welfare assessment will be aggregated at a colony level to compare welfare between colonies with more and less access to food.

Why we funded this project

We funded this project because it seeks to evaluate welfare impacts of population density — likely a common cause of welfare issues — in a relatively abundant species, using a robust approach that is only possible thanks to the species-specific knowledge of the PI and her collaborators. For example, we value their use of image analyses to identify haul-out site suitability and welfare impacts on a fine spatial scale that takes into account both environmental and social factors. We also valued the focus on juvenile welfare and the conflict of interest that may arise between parent and offspring. Improving our understanding of the fundamental relationships between population density, parental care, and juvenile welfare will allow the results of this project to extend to other contexts.


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