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.
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.
Pre- and Post-release Welfare Indicators for Recovery Programs
Grantee: Laney Hayward Nute
Institution: University of Mississippi
Project summary
Captive-raised animals translocated to the wild often suffer high mortality. This project seeks to identify a set of pre- and post-release indicators of welfare to target released Attwater’s prairie chickens for management interventions. Pre-release, the coping styles of birds will be identified during routine cage transfers and handling. Following release, the fine-scale movements of released birds will be traced using customized data loggers. Investigating the association of pre- and post-release behaviors with body condition, daily environment changes, and survival, will identify which birds from future releases will need special attention to aid their survival. The researchers hypothesize that individuals with a more proactive personality will disperse further post-release and suffer higher mortality rates compared to more reactive individuals.
Grantee: Laney Hayward Nute
Institution: University of Mississippi, United States
Grant amount: $30,000
Grant type: Small grants
Focal species: Attwater's Prairie-chicken (Tympanuchus cupido attwateri)
Conservation status: Endangered
Disciplines: Wildlife rehabilitation, animal behavior, ornithology
Research location: United States
Project summary
Captive-raised animals translocated to the wild often suffer high mortality; for example, Attwater’s prairie chickens have had a survival rate of only 17-18% post-release in recent years. The focus of many translocation programs is on investing resources to produce more individuals rather than to improve the welfare of released animals. This project seeks to reduce the stress and suffering of released Attwater's prairie chicken by identifying a set of pre- and post-release indicators of welfare to target individuals for management interventions. Pre-release, the coping styles of individual birds will be identified during routine cage transfers and handling. Following release, the fine-scale movements of released birds will be traced using customized data loggers. By investigating the association of pre- and post-release behaviors with the body condition, daily environment changes, and ultimate survival of these individuals, it will be possible to identify which birds from future releases will need special attention to aid their survival. The researchers hypothesize that individuals with a more proactive (“bold”) personality will disperse further post-release and suffer higher mortality rates compared to more reactive individuals.
Why we funded this project
Understanding of associations between coping style/personality and welfare outcomes/challenges could be used to predict at-risk individuals in need of interventions such as supplemental feeding, translocation, or medical treatment. For wildlife rehabilitation or “headstarting” programs, coping style associations could also be used to inform rearing protocols to promote resilient styles that result in improved individual welfare in the wild.
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Developing a consensus profile of wild animal welfare: integrating non-invasive monitoring of the gut microbiome with stress physiology and behavior
Grantee: Sam Sonnega
Institution: University of Massachusetts Dartmouth
Project summary
This project will characterize the gut microbiome of white-footed mice and investigate its relationship with their stress physiology and behavior. Mice will be trapped and fecal samples collected to measure glucocorticoid concentrations and gut microbiome composition. Concurrently, open-field trials will be conducted to assess individual variation in cognitive bias. Perception of predation risk will also be experimentally manipulated by exposing mice to playbacks of predator noises. By correlating the gut microbiome with behavioral and endocrine metrics, a consensus profile of the mice’s welfare will be developed that reflects the complexity of their responses to environmental perturbations, and how those responses can scale up to population and ecosystem level changes via the demographic effects of stress in a “landscape of fear.”
Grantee: Sam Sonnega
Institution: University of Massachusetts Dartmouth, United States
Grant amount: $29,130
Grant type: Small grants
Focal species: White-footed mice (Peromyscus leucopus)
Conservation status: Least concern
Disciplines: Physiology, animal welfare science, animal behavior
Research location: United States
Publications
Sonnega, S. and Sheriff, M.J. (2024). Harnessing the gut microbiome: a potential biomarker for wild animal welfare. Frontiers in Veterinary Science, 11. https://doi.org/10.3389/fvets.2024.1474028
Project summary
This project will characterize the gut microbiome of wild white-footed mice (Peromyscus leucopus) and investigate its relationship with their stress physiology and behavior, including how it changes in response to ecological pressures. While a growing literature from lab-based studies has demonstrated the link between the gut microbiome and regulation of host physiology and behavior, the generality of these findings in ecological contexts remains largely untested. As part of this project, mice will be trapped during different seasons and fecal samples will be collected from which to measure both glucocorticoid concentrations and gut microbiome composition. Concurrently, open-field trials will be conducted to assess individual variation in cognitive bias towards optimism or pessimism (a well-established behavioral indicator of affective state). The mice’s perception of predation risk will also be experimentally manipulated by exposing free-living mice to playbacks of predator noises. By correlating the gut microbiome with both behavioral and endocrine metrics, a consensus profile of the mice’s welfare will be developed that reflects the complexity of their responses to both predictable and unpredictable environmental perturbations, and how those responses can scale up to population and ecosystem level changes via the demographic effects of stress in a so-called “landscape of fear.”
Why we funded this project
We are interested in the development and validation of the gut microbiome as a welfare indicator. Every additional indicator strengthens the interpretation of others, but understanding the gut microbiome may be especially important because it is part of the causal chain linking what is going on in the animal’s brain to fecal metabolites, which are often analyzed as a non-invasive and time-integrated record of physiological stress. This project also links and builds on other gut microbiome research we have funded (by Melissa Bateson, Pablo Capilla-Lasheras, and Davide Dominoni).
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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.
The animal welfare of animal warfare: how inter-group interactions affect wild animal wellbeing
Grantee: Dominic Cram
Institution: University of Cambridge
Project summary
This project will investigate the welfare consequences of inter-group conflicts in wild Kalahari meerkats. It will use an established dataset of meerkat behavior, body weight, territory use, and reproduction, covering more than 500 inter-group interactions over 14 years. Identifying the short- and long-term welfare implications of inter-group interactions in meerkats will shed light on similar animal warfare in group-living insects, fish, birds, and mammals. Clarifying the circumstances that lead to the most harmful battles will provide a first step in understanding how their frequency could be reduced, which could limit the suffering they cause and enhance the welfare of group-living wild animals.
Grantee: Dominic Cram
Institution: University of Cambridge, United Kingdom
Grant amount: $29,965
Grant type: Small grants
Focal species: Meerkats (Suricata suricatta)
Conservation status: Least concern
Disciplines: Animal behavior, population ecology, community ecology
Research location: United Kingdom, South Africa
Project summary
Fierce group conflicts are not uniquely human, and many group-living animals regularly engage in “animal warfare.” These inter-group interactions play an influential role in natural population regulation, yet the health and well-being consequences for those involved remain unclear. Conservation and management interventions are currently developed with little understanding of how large-scale conflict affects welfare in wild animals. Given that anthropogenic habitat loss and climate change could increase the frequency of inter-group battles, there is an urgent need to investigate the welfare cost of animal warfare. This research program will investigate the welfare consequences of intergroup interactions and fights in wild Kalahari meerkats (Suricata suricatta). The project will take advantage of an established dataset of meerkat behavior, body weight, territory use, and reproduction, covering more than 500 inter-group interactions over 14 years. Identifying the short- and long-term welfare implications of inter-group interactions in meerkats will shed light on similar animal warfare in group-living insects, fish, birds, and mammals. Clarifying the circumstances that lead to the most harmful battles will provide a first step in understanding how their frequency could be reduced, which could limit the suffering they cause and enhance the welfare of group-living wild animals.
Why we funded this project
This project addresses a neglected topic related to our research priority of understanding conflicts of interest between wild animal groups. We especially appreciate this project’s holistic approach to welfare assessment, its attention to indirect effects (collateral damage) of animal conflicts in the form of costs to orphaned juveniles and the creation of a “landscape of fear,” and the openness of the investigators to considering interventions that could elevate wild animal welfare above its natural baseline.
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Development of octopus mind in the wild: a behavioral, ecological and evolutionary investigation into sentience and emotional states in Octopus insularis juveniles
Grantee: Michaella Andrade
Institution: Federal University of ABC
Project summary
There is evidence that evolutionary pressures can cause behaviors with opposite meanings to develop opposite forms, the way a frown is the opposite of a smile. One way to understand the expression of emotion in animals may therefore be to identify pairs of behaviors that are opposites. In octopuses, which are increasingly being recognized as sentient, colors can be signals of emotional valence during conflict and other situations. Yet no study has tested whether octopuses have opposite pairs of color signals. This project will produce descriptions of evolutionary and behavioral patterns that reflect the emotional states and sentience of juvenile octopuses, which may contribute to the welfare of octopuses and other invertebrates.
Grantee: Michaella Andrade
Institution: Federal University of ABC, Brazil
Grant amount: $37,959
Grant type: Challenge grants
Focal species: Octopuses (Octopoda sp.)
Conservation status: Data deficient
Disciplines: Sentience, animal behavior
Research location: Brazil
Publications
Andrade M.P., et al. (2023). Assessing Negative Welfare Measures for Wild Invertebrates: The Case for Octopuses. Animals, 13(19), 3021. https://doi.org/10.3390/ani13193021
Project summary
Welfare refers to the quality of life of animals that possess sentient capacity and emotional states. Although the precursors of sentience were possibly present on the planet as early as 890 million years ago, the evolution of sentience is still poorly understood. However, cephalopods are increasingly being recognized as sentient, yet we do not know about this phenomenon in juvenile wild animals. In this sense, studies with wild animals can be beneficial for finding a wider range of ecological triggers and their relationship with behaviors.
There’s evidence that evolutionary pressures can cause behaviors with opposite meanings to eventually develop opposite forms, the way a frown is the opposite of a smile. One way to understand the expression of emotion in animals therefore may be to find pairs of behaviors that are opposites. In octopus, colors can be a signal of emotional valence during conflict and other contexts. Although researchers began to see this dimension in octopuses, no study has tested whether opposite pairs of color signals are present in octopuses. This project will produce descriptions of evolutionary and behavioral patterns that reflect the emotional states and sentience for juvenile octopuses, which may contribute to the welfare of octopuses and other invertebrates.
Why we funded this project
Although octopuses are widely assumed to be sentient at the adult stage, no studies that we are aware of have examined sentience at earlier life stages. As the vast majority of octopuses alive at any one time are juveniles, and octopuses have enormously high juvenile mortality, the question of when in their development sentience arises is particularly important. This project is also interesting because it will teach us about what the lives of juvenile octopuses are like and the extent to which welfare effects are mediated by personality traits.