MSc by Research in Psychology (Animal Behaviour)

Are you interested in studying for an MSc by Research at the Centre for Research in Animal Behaviour (CRAB)? This 2-year degree offers you the opportunity to conduct independent research and receive in-depth research training to support your progression into various career paths or continuing into a PhD.

How to apply

Prospective students should contact the relevant academic staff in the first instance to discuss possible research areas and their application. Applicants have to prepare a research proposal which must be agreed with the supervisor before applying through the online application portal (see link below). The application process involves a formal review of the proposal and interview with a selection panel led by the Director of Postgraduate Research of the Department of Psychology.

Entry requirements: Applicants should have obtained (or are about to obtain) a 2.1 or above in Psychology, Biology or a related undergraduate degree. If you have a 2.2 degree but also obtained a Masters degree, you are also eligible. For international applicants, English requirements are IELTS (or equivalent) overall score 6.5 (no less than 6.0 in any section).

More information can be found here:

Training opportunities

The MSc by Research is a 2-year research degree programme (or 3 years part time) and students do not complete taught components. However, there are numerous opportunities to develop skills and experience. We have weekly meetings (CRAB lunches) to discuss papers, recent project progress or meet with visitors. We have regular external speakers giving seminars in the Animal Behaviour Research Seminar Series, as well as in the wider Department and College. The University's Researcher Development Programme offers an extensive choice of training and development events exclusively designed for our postgraduate research students.


You will be assessed by a written thesis or scientific paper of up to 40,000 words in length that is externally examined.


Below are some opportunities to join research projects that are running in CRAB as an MSc-by-Research student. Please note that this is not an exhaustive list but is intended to provide an overview of the types of projects that are possible. We strongly encourage students to approach staff with ideas for their own project or ideas to extend the projects outlined below.

Social Dynamics in Captive Animals: Implications for Welfare and Productivity
Advancing our understanding of social interactions in food production animals is of fundamental importance for maximizing productivity and welfare. Previous research in this area has tended to focus on how group attributes such as group size, density and composition impact on welfare and productivity. Currently however, we know very little about how social relationships form within groups and how the structure and function of these relationships change across development, environments and contexts. This gap in knowledge is a major issue in industries such as the dairy industry where social stress that results from social disruption (due to the addition and removal of individuals) can have significant economic and welfare implications. For example, the perturbation of group structure has management implications and can result in increased aggression and stress and reduced feed intake and milk yield. There are many areas for possible projects around this topic. For example, one approach could be to study the temporal structure of social relationships in farmed animals across development, environments and contexts with the aim of improving the management of social relationships.
Relevant papers
  • Boyland et al. (2016) "The social network structure of a dynamic group of dairy cows: From individual to group level patterns." Applied Animal Behaviour Science 174: 1-10.
  • Diosdado et al. (2015) "Classification of behaviour in housed dairy cows using an accelerometer-based activity monitoring system." Animal Biotelemetry 3.1: 1.
  • Rose & Croft (2015) "The potential of Social Network Analysis as a tool for the management of zoo animals." Animal Welfare 24.2: 123-138.
For more information about this project please contact Prof. Darren Croft


The Evolution of Social Relationships
In many animals, affiliative relationships between pairs of individuals are a pervasive part of the social landscape. In recent years, scientists have shown that these bonds are associated with fitness and have clear associations with specific neural mechanisms and neuroendocrine systems. Yet there remains a great deal of research to be done and possibilities for projects on this topic are numerous. For example, there are considerable gaps in our understanding of the strength and stability of social relationships and their associations with fitness, especially amongst members of the dispersing sex. We are also only just beginning to understand the heritable (genetic) basis of differences in social tendencies between group mates. There are also large gaps in our appreciation of the importance of the different types of social ties in polyadic networks (ie. direct vs. indirect connections). Access to a large dataset comprised of social network information on hundreds of free-ranging rhesus macaques could form the basis of student-led projects in this area.

Relevant papers

  • Silk et al. (2003) "Social bonds of female baboons enhance infant survival." Science 302: 1231-1234.
  • Brent et al. (2013) “Genetic origins of social networks in rhesus macaques.” Scientific Reports 3, 1042. DOI 10.1038/srep01042.
  • Brent et al. (2014) “The neuroethology of friendship.” Annals of the New York Academy of Sciences. DOI 10.1111/nyas.12315.
  • Brent (2015) “Friends of friends: Are indirect connections in social networks important to animal behaviour?” Animal Behaviour 103: 211-222. doi:10.1016/j.anbehav.2015.01.020.
For more information about this project please contact Dr Lauren Brent
Collective or Coordinated Vigilance in Flocks of Pheasants

Animals living in groups may gain benefits from the vigilance behaviour of groupmates. Individuals can forgo costly vigilance behaviour and instead spend time foraging, with the assurance that if a predator approaches their groupmates will alert them. However, with each group member wanting to selfishly avoid costly vigilance, it is unclear exactly how each individual decides whether to engage in guarding and consequently what group-level patterns of vigilance emerge. Two extreme patterns may be expected. First, all individuals may simultaneously be vigilant when threat levels are high and conversely none are vigilant when expectations of predation are low. This leads to very high collective vigilance in risky situations, but low or no vigilance when no predator is expected. Second, a minimal number of individuals are vigilant at all times and this role is alternated between group members. This leads to continuous lower level vigilant coverage regardless of predation risk.


Pheasants are gregarious during the breeding season with a territory-holding male hosting a harem of females. Individual’s vigilance levels vary depending on harem size (Whiteside et al. 2016). This project will extend this work to explore how individuals adjust their vigilance behaviour according to a) the vigilance of others in the group; b) the threat level by predators. You will use detailed observations coupled with automated behavioural measurements using accelerometers to generate patterns of vigilance exhibited by groups of captive pheasants. You will manipulate group compositions, individuals’ motivation to be vigilant and threat levels and explore how these alter vigilance patterns.

Relevant papers

  • Whiteside et al. (2016) “Males and females differentially adjust vigilance levels as group size increases: effect on optimal group size.” Animal Behaviour 118: 11-18.
Fieldwork will take place between February and June in Mid Devon. Accommodation is available on site. This project forms part of a larger ERC-funded study by Joah Madden looking at the fitness consequences of differential cognitive performance in pheasants. As such, you will join the established Pheasant Ecology and Cognition group (Twitter: @pec_exeter). For more information about this project please contact Dr Joah Madden

Grey squirrels and pine martens: the role of stress in suppressing grey squirrel populations

This project aims to investigate stress as a mechanism for the decline in grey squirrel numbers by testing the hypothesis that pine marten presence is associated with behavioural and physiological indicators of stress in grey squirrels 

The project will make use of grey squirrels from UK culling programmes, comparing populations where pine martens are either present or absent. Videos from feeders will be examined to measure behavioural indicators of fear, such as increased vigilance and decreased feeding activity. Faecal samples will be collected to test for stress hormones, which indicate acute stress. Body condition will be used to test if the presence of pine martens has resulted in decreased health. To determine if grey squirrels are experiencing chronic stress in the presence of pine martens the squirrels’ brains will also be collected in order to measure whether they show stress-related suppression of adult hippocampal neurogenesis. This is a new measure used mainly in the fields of psychiatry and animal welfare and will show whether there are cumulative stress effects of co-existing with pine martens over a grey squirrel’s lifetime. 

Together, these data will provide a key indicator of whether stress-mediated indirect effects are the cause for the grey squirrel decline in the presence of pine martens. Pinpointing the mechanism affecting interactions between these two species will help to better manage grey squirrel invasion throughout Europe and provide a novel approach for determining indirect effects in native/invasive species interactions.

For more information about this project please contact Dr Lisa Leaver


Navigation and Foraging of Bees in the Natural Landscape

Navigation in bees is influenced by landmarks, and in heterogeneous fragmented landscapes with major landmark structures, those can significantly determine where and how bees forage, subsequently affecting pollination services. In this project we will investigate how bees adjust their behaviour to such conditions. In this project work involves field observations of foraging bees in South Devon and implementation of the data into predictive models of distribution of pollination services. The South Devon Area of Outstanding Natural Beauty (AONB), offers convenient field sites. Many small and diverse agricultural holdings are separated by wide large-sized high-hedges grown on earth banks, the traditional Devon hedges. It is also a hotspot of biodiverse habitats many of which have been recently mapped out in the B(iodiversity)-Lines project which will connect them with corridors to conserve biodiversity and oppose the decline in bees, the major group of pollinators. The data collected in this project, scientific conclusions and outcomes of model testing will feed directly into the management of the AONB landscape to enable the assessment of the effectiveness of B-Lines and landscape management practices on the pollination services provided by bees. This project offers multidisciplinary academic training in the behavioural and ecological sciences and opportunities for non-academic training in landscape management and the development and implementation of environmental policies. 


Relevant papers

  • Hempel de Ibarra et al. (2009). “Preferred viewing directions of bumblebees (Bombus terrestris L.) when learning and approaching their nest site.” Journal of Experimental Biology 212: 3193-3204.
  • Collett et al. (2006). “Navigational memories in ants and bees: Memory retrieval when selecting and following routes.” Advances in the Study of Behaviour 36: 123-174.
  • Osborne et al. (2008) “Bumblebee flight distances in relation to the forage landscape.” Journal of Animal Ecology 77: 406-415.
  • Stefan-Dewentter & Kuhn (2003) “Honeybee foraging in differentially structured landscapes.” Proceedings of the Royal Society B 270: 569-575.
  • Becher et al. (2014) “BEEHAVE: A systems model of honeybee colony dynamics and foraging to explore multifactorial causes of colony failure.”  Journal of Applied Ecology 51: 470-482.

For more information about this project please contact Dr Natalie Hempel

Specialisation in Cooperating Groups of Animals

Animals that cooperate in groups often show specialisation, whereas in closely-related species all individuals appear to be identical. The reasons for these differences are not well understood. In this project we will perform a comprehensive survey of the scientific and natural history literature to record differentiation of workers in colonies of bees, ants and wasps. We will record the size variation and different morphologies of workers, as well as known differences in behavioural tendencies in workers. We will collect data on the environment, ecology and colony life history of species, such as seasonality, colony size, reproductive strategy and food supply characteristics and the incidence of predation on nests, many of which are well known for many species, which will enable us to understand what aspects of colony life lead to differentiation. This project offers training in phylogenetically-controlled analysis of large data sets.

Relevant papers

  • Ferguson-Gow et al. (2014) “Colony size predicts division of labour in attine ants.” Proceedings of the Royal Society B 281: 20141411. doi: 10.1098/rspb.2014.1411.
  • Higginson et al. (2012) “Masquerade is associated with polyphagy and larval overwintering in the Lepidoptera.” Biological Journal of the Linnean Society 106: 90-103.
For more information about this project please contact Dr Andy Higginson

The Evolution of Behavioural Courtship Displays

The extensive body of theory on sexual selection explains how conspicuous morphological ‘ornaments’, such as bright colour patches and exaggerated tail feathers, can be stably maintained by female preferences despite their clear survival costs. However, this theory does not readily apply to the flexible behavioural components (e.g. energetic movement patterns, courtship song) that are a prominent feature of many displays. Such responsive, dynamic courtship elements raise important questions regarding the strategic allocation of male courtship effort and how honest signalling of quality can be maintained when courtship intensity is flexible. To address these issues, this project will use evolutionary models and computer simulations to understand how male strategies for allocating courtship effort coevolve with static morphological ornaments and female preferences for those different display components.

Relevant papers 

  • Barske et al. (2011) “Female choice for male motor skills.” Proceedings of the Royal Society B 278: 3523-3528.
  • Kuijper et al. (2011) "A guide to sexual selection theory." Annual Review of Ecology, Evolution, & Systematics 43: 287-312.
  • South et al. (2012) “Female preference for male courtship can drive the evolution of male mate choice.” Evolution 66: 3722-3735.
For more information about this project please contact Dr Tim Fawcett

The Social Consequences of Sexual Harassment in a NonHuman Primate

Male sexual coercion of females is a direct result of an evolutionary conflict of interest between the sexes over reproduction. When males sexually harass or coerce females into mating they are actively increasing the mating rate towards their own optimum by increasing the number and / or quality of females whose eggs they fertilize. Work on sexual conflict has focused primarily on sexually antagonistic traits that are energetically costly to females, that physically damage females, that directly inhibit female foraging or directly increase female exposure to predators. More recently however, work has demonstrated that the presence of sexually harassing males can alter female social behaviour in a way that is likely to decrease the probability of females engaging in positive social associations with others in the population; that is, females can experience a social cost. This project aims to examine the social consequences of female sexual harassment by males in free-roaming groups of rhesus macaques (Macaca mulatta) on the island of Cayo Santiago, Puerto Rico. Potential questions that could be addressed include ‘does being a target of sexual harassment correlate with social performance?’, ‘do experiences with sexual harassment predict changes in social network structure?’ and ‘does target social rank matter in the prevalence and consequences of sexual harassment?’

Relevant papers

  • Chapman et al. (2003) “Sexual conflict.” Trends in Ecology & Evolution 18: 41-47.
  • Darden et al. (2009) “Social implications of the battle of the sexes: sexual harassment disrupts female sociality and social recognition.” Proceedings of the Royal Society B 276: 2651-2656.
  • Darden & Watts (2012) “Male sexual harassment alters female social behaviour towards other females.” Biology Letters 8: 186-188.
  • Parker (2006) “Sexual conflict over mating and fertilization: an overview”. Philosophical Transactions of the Royal Society B 361: 235-259.
  • Silk (2007) “Social components of fitness in primate groups” Science 317: 1347-1351.
For more information about this project please contact Dr Safi Darden or Dr Lauren Brent

Executive control and investment decisions in squirrels: an ecological perspective on planning for the future

An individual’s ability to inhibit a pre-potent response or “executive control”, often conceptualised as self-control, is an important aspect of cognitive flexibility and has been used as an assay of cognitive performance in a number of studies. Psychological studies of executive control often involve abstract tests in the laboratory to measure specific inhibition-of-response times across individuals. Executive control has been identified as an important factor in gambling and drug addiction, but the importance of executive control in less extreme behaviours has been largely overlooked. One such behaviour that is crucial for wellbeing and ensuring future security is investment behaviour, which necessarily entails planning for the future. Investment in the future can take many forms across species, and requires some degree of executive control because an individual needs to inhibit current use of an asset to store it away for a future which is, inevitably, uncertain. Squirrels are an ideal model species for studying the link between executive control and investment because they are natural and prolific investors. Squirrels are well known to cache food widely for later use, a risk-averse behavioural strategy that involves storing nuts during times of plenty in order to provide a reliable source of food when resources are scarce. Squirrels not only decide whether to cache a food item, eat it, or reject it, but caching itself involves a series of predation and pilferage risk trade-offs, which reliably co-vary with food value. As such, investment decisions and effort directly reflect future discounting and ought to be linked to executive control. The MRes project student will investigate the role of executive control in investment decisions by wild squirrels. He/she will collect experimental and observational data on squirrels, get involved in devising field experiments to verify abstract laboratory tests of executive control, and measure  investment outcomes, a real-life measure of delaying gratification. This project could be carried out on red or grey squirrels (Sciurus vulgaris and S. carolinensis).


Relevant papers

  • Vernouillet et al. (2016). “Inhibition in Clark’s nutcrackers (Nucifraga columbiana): results of a detour-reaching test.” Animal Cognition 19:661–665. DOI 10.1007/s10071-016-0952-y
  • McLean et al. (2014) “The evolution of self control”. Proceedings of the National Academy of Sciences USA 111: E2140–E2148.

For more information about this project please contact Dr Lisa Leaver

Male African Elephant Society

A comprehensive understanding of male elephant ecological and social requirements is vital to effectively target conservation efforts, particularly in light of ever-increasing anthropogenic and environmental change.


Research in the Boteti River region of the Makgadikgadi Pans National Park of Botswana, an area dominated by male elephants, is giving us a rare opportunity to unravel more about the nature and significance of male elephant social behaviour. Male elephant social systems are complex. Males of all ages prefer to be closest to the old bulls in social groups and it is likely that they, like female matriarchs, are the repositories of social and ecological knowledge that is vital to the stable functioning of populations. Understanding how these bulls can influence key behaviours such as navigation, foraging and dominance in behaviours, including crop raiding will help shape future conservation efforts that ensure that their social, alongside their ecological needs, are taken into account.


There are many areas for possible projects around the social behaviour of male elephants, including how they communicate with each other and potentially pass on information (e.g. temporal gland secretions, tactile communications and, vocalisations) and by what means are males accessing each other.

Relevant papers

For more information about this project please contact Prof. Darren Croft

The Psychology of Foraging in Bees

Foraging theory has been moving beyond optimal foraging models to incorporate limits on perfect behaviour due to psychological, physiological and informational constraints. Individuals of species interacting with others, which underpin ecosystem function, are usually assumed not to be variable, but this is unrealistic with variation due to differences in learning and experiences. This project will focus on laboratory studies of social bumblebees foraging on artificial flowers in order to understand how bees learn, why they are variable in their behaviour, and what this means for ecosystem services and stability.

Relevant papers

  • Fawcett et al. (2015) “The evolution of mechanisms underlying behaviour.” Current Zoology 61: 221-225.
  • Higginson & Houston (2015) “The influence of the food-predation trade-off on the foraging behaviour of central-place foragers.” Behavioral Ecology & Sociobiology 69: 551-561.
For more information about this project please contact Dr Andy Higginson

Winner and Loser Effects in Sport

Winner and loser effects, in which the experience of winning or losing a contest affects the ability to win future contests, are widespread across the animal kingdom. Experiences of victory and defeat precipitate a range of physiological and behavioural changes that make winners more likely to win again and losers to carry on losing. Recently, evolutionary biologists have argued that similar effects exist in human sport and may underlie winning and losing ‘streaks’, but this idea has not yet been rigorously tested. This project will address that gap, by using extensive data sets from sports websites to quantify the factors affecting changes in performance across an athlete’s career. Advanced statistical techniques, based on generalised linear mixed models (GLMMs), will be used to disentangle the direct psychological impact of winning and losing from other fluctuations in form, guided by testable predictions from evolutionary models of contest behaviour.


Possible research questions include the following:

  • How does the impact of winning and losing change with age?
  • Do male and females respond differently to victories and defeats?
  • Does the magnitude of a win/loss alter the impact of winner/loser effects?

Relevant papers

  • Booth et al. (1989) “Testosterone, and winning and losing in human competition.” Hormones & Behavior 23: 556-571.
  • Fawcett & Johnstone (2010) “Learning your own strength: winner and loser effects should change with age and experience.” Proceedings of the Royal Society B 277: 1427-1434.
  • Dugatkin & Reeve (2014) “Winning, losing, and reaching out.” Behavioral Ecology 25: 675-679.
For more information about this project please contact Dr Tim Fawcett

Living Outside of Groups: Pathology, Social Exclusion, or Adaptive Strategy?

In humans, there is a subset of people who are inherently asocial. These socially isolated people suffer considerable health consequences and have reduced survival rates. This situation may be maladaptive, caused by neurocognitive pathologies or by competitive exclusion by group members, or may represent an adaptive strategy. In nonhuman primates, where we are better able to address questions about adaptive benefits, asocial individuals have also been documented, including individuals who fail to gain permanent access to a social group. However, most field studies insert researchers into social groups and as a result collect data on individuals who are somewhat socially successful.  The free-ranging rhesus macaques (Macaca mulatta) of the Cayo Santiago field station live on island and so individuals rarely go unaccounted for, allowing us to collect data on both group-living and ‘asocial’ animals. There are many possible projects on the causes and consequences of asociality in this system. For example, one approach could be to compare the behaviours of asocial and group-living individuals, such as their tendencies to be aggressive toward others, or to engage in risk-taking behaviours. One could also investigate the survival and health consequences of these different strategies.

Relevant papers

  • Holt-Lunstad et al. (2010) “Social relationships and mortality risk: a meta-analytic review.” PLoS Medicine 7: e1000316.
  • Silk (2014) “The adaptive value of sociality.” In Mitani et al. (eds) The Evolution of Primate Societies (Univ. Chicago Press, Chicago, IL).
  • Brent et al. (2013) “Genetic origins of social networks in rhesus macaques.” Scientific Reports 3, 1042. DOI 10.1038/srep01042.
  • Brent et al. (2014) “The neuroethology of friendship.” Annals of the New York Academy of Sciences. DOI 10.1111/nyas.12315.

For more information about this project please contact Dr Lauren Brent

Learning to Be Ill: An Evolutionary Approach to Understanding Mental Disorders

Why we are disposed to mental illnesses is not well understood. This project will develop an emerging paradigm in modelling mental illness founded in evolutionary ecology. We will develop computer models of learning and decision-making that capture the phenomena seen in depression, anxiety, PTSD and/or OCD. The insights from the models will improve treatments for these conditions. This project offers training in evolutionary modelling and computer programming.

Relevant papers 

  • Trimmer et al. (2015) “Adaptive learning can result in a failure to profit from good conditions: implications for understanding depression.” Evolution, Medicine & Public Health 2015:123-135.
For more information about this project please contact Dr Andy Higginson

The Evolution of Individual Recognition

Few animals are known to recognise members of their own species as individuals, i.e. to learn and recall unique individuals during subsequent encounters. Yet this ability is a prerequisite to engaging in a host of complex social strategies. A clearer understanding of the evolution of individual recognition is thus an important component of our understanding of the evolution sociality. Students wishing to work in this area would be invited to explore the evolutionary history of individual recognition using modern comparative analyses.

Relevant papers 

  • Tibbetts & Dale (2007) "Individual recognition: it is good to be different." Trends in Ecology & Evolution 10: 529-537.
  • Brent et al. (2014) “The neuroethology of friendship.” Annals of the New York Academy of Sciences. DOI 10.1111/nyas.12315.
  • Seyfarth & Cheney (2015) "The evolution of concepts about agents: or, what do animals recognize when they recognize an individual?" The Conceptual Mind: New Directions in the Study of Concepts: 57.
For more information about this project please contact Dr Lauren Brent

Understanding Obesity by a Comparative Approach

Evolutionary approaches to understanding fattening in animals offer potential solutions to the obesity epidemic. Often models are constructed based on assumptions about environments in which animals, including humans, evolved. However, there is often little evidence for these assumptions. In this project we will build and analyse a database of the ecology and life history of species for which the average level of fat storage is known. This will involve collecting data from the scientific and natural history literature on variables such as seasonality, body size, reproductive strategy and food supply characteristics. This project offers training in phylogenetically-controlled analysis of large data sets.

Relevant papers 

  • Higginson et al. (2016) “Fatness and fitness: Exposing the logic of evolutionary explanations for obesity.” Proceedings of the Royal Society B 283: 20152443. doi:10.1098/rspb.2015.2443
  • McNamara et al. (2016) “The influence of the starvation-predation trade-off on the relationship between ambient temperature and body size among endotherms.” Journal of Biogeography 43: 809-819. doi:10.1111/jbi.12695 
For more information about this project please contact Dr Andy Higginson

 Other projects: