MSc(Res) Behaviour, Ecology and Evolution

The Master of Science by Research degree in Behaviour, Ecology and Evolution is a 12-month, research only degree, in which the candidate will undertake a supervised research project in the broad area of Behaviour, Ecology and Evolution, in the School of Biology, University of St Andrews.

The candidate will be based in the interdisciplinary Centre for Biological Diversity (CBD), based in the centre of St Andrews. The CBD links researchers in evolution, behaviour, ecology, molecular biology and biodiversity, plus researchers in other Schools across St Andrews. Research themes include: the mechanistic causes and the ecological and evolutionary consequences of animal behaviour, with strengths in behavioural ecology, animal cognition, social evolution and social learning; evolutionary and population genetics, including the genetic basis of population divergence and speciation; animal-plant interactions, including pollinator biology; conservation biology, focusing in particular on the measurement of broad-scale patterns of biodiversity and biodiversity change. These themes are underpinned and guided by theoretical evolutionary ecologists and geneticists, asking fundamental questions about the causes and consequences of organismal interaction. Our final objective is to advance this scientific understanding of the diversity of life to contribute pro-actively to policy that helps protect and nurture biological diversity.

Candidates may approach potential supervisors in the CBD directly or via advertised projects listed below.


Evolution of Same-Sex Sexual Behaviour

Supervisor: Dr Nathan Bailey

Research area (s): Evolution of Same-Sex Sexual Behaviour

Research description: Same-sex sexual behaviour (SSB) is frequently portrayed as an evolutionary paradox. But is it? The Bailey lab has explored causes of its expression at low but persistent levels in insects. This project will apply a comparative approach to test hypotheses about the evolution of SSB, capitalising on an extensive resource of videotaped behavioural interactions across multiple field cricket species. The student will have flexibility to tailor the project to their specific interests. The project is based at St Andrews, in collaboration with the co-supervisor Dr. Kevin Judge at MacEwan University.

Relevant references:

(1) Bailey NW, Zuk M (2009) Same-sex sexual behavior and evolution. Trends in Ecology and Evolution 24:439-446.

(2) Hoskins JL, Ritchie MG, Bailey NW (2015) A test of genetic models for the evolutionary maintenance of same-sex sexual behaviour. Proceedings of the Royal Society of London, B 282:20150429.

(3) Judge KA, Ting JJ, Schneider J, Fitzpatrick MJ (2010) A lover, not a fighter: mating causes male crickets to lose fights. Behavioral Ecology and Sociobiology 64:1971-1979.

(4) Bailey NW, Hoskins JL, Green J, Ritchie MG (2013) Measuring same-sex sexual behaviour: the influence of the male social environment. Animal Behaviour 86:91-100.

(5) Bailey NW, French N (2012) Same-sex sexual behaviour and mistaken identity in male field crickets (Teleogryllus oceanicus). Animal Behaviour 84:1031-1038.

Subject area(s): Animal Behaviour, Behavioural Ecology, Evolutionary Biology, Comparative Analysis

Keywords: aggression, animal weaponry, field crickets, mating systems, same-sex sexual behaviour, sexual conflict, sexual dimorphism, sexual selection

Natural and Sexual Selection on Insect Signals in Oceania

Supervisor: Dr Nathan Bailey

Co Supervisor: Prof Leigh Simmons, University of Western Australia

Research area (s): Evolution and Animal Behaviour

Research description: How do animal signals evolve under conflicting evolutionary pressures? Sexual selection favouring conspicuous signals to attract mates can be counteracted by natural selection selection acting against energetic or predation costs. This project will use field crickets to examine how variable climatic factors constrain, or alternatively facilitate, the evolution of chemical signals used during mate choice in populations across Australia and Oceania. The student will have flexibility to shape the project to suit their own interests. The project is based in St Andrews, in collaboration with the co-supervisor Prof Leigh Simmons at the University of Western Australia.

Relevant references:

(1) Pascoal S, Risse JE, Zhang X, Blaxter M, Cezard T, Challis RJ, Gharbi K, Hunt J, Kumar S, Langan E, Liu X, Rayner JG, Ritchie MG, Snoek BL, Trivedi U, Bailey NW (2020) Field cricket genome reveals the footprint of recent, abrupt adaptation in the wild. Evolution Letters.

(2) Berson JD, Zuk M, Simmons LW (2019) Natural and sexual selection on cuticular hydrocarbons: a quantitative genetic analysis. Proceedings of the Royal Society of London, B. 286:20190677.

(3) Moran PA, Hunt J, Mitchell C, Ritchie MG, Bailey NW (2019) Behavioural mechanisms of sexual isolation involving multiple modalities and their inheritance. Journal of Evolutionary Biology. 32:243-258.

(4) Berson JD, Simmons LW (2019) Female cuticular hydrocarbons can signal indirect fecundity benefits in an insect. Evolution. 73:982-989.

(5) Pascoal S, Mendrok M, Wilson AJ, Hunt J, Bailey NW (2017) Sexual selection and population divergence II. Divergence in different sexual traits and signal modalities in field crickets (Teleogryllus oceanicus). Evolution. 71:1614-1626.

Subject area(s): Animal Behaviour, Behavioural Ecology, Evolutionary Biology, Evolutionary Ecology

Keywords: animal signal, biogeography chemical ecology, cuticular hydrocarbon, field crickets, mate choice, natural selection, sexual selection

Evolutionary developmental biology

Supervisor: Dr David E. K. Ferrier

Research area(s): Evolutionary developmental biology

Research description: We seek to understand how the diversity of animal forms have evolved via changes to their development, usually taking the homeobox genes of the Hox/ParaHox and related clusters as a starting point. A variety of invertebrate species are studied (including amphioxus, Ciona, annelids, arthropods, cnidarians and sponges), with the aim of focusing on major transitions in animal evolution, including the origins of the animal kingdom, the origin of the bilaterally symmetrical animals (bilaterians) and the origin of chordates and vertebrates.

Relevant references: See

Subject area(s): Evolutionary biology, Developmental Biology

Keywords: Evo-devo, regeneration, embryology, genomics

Animal cognition

Supervisor: Dr Sue Healy

Research area (s): Animal cognition

Research description: Cognition plays a central role in the lives of many animals, whether with regard to learning and remembering where to find food or home, making decisions over choices of food or mates or in interacting with others.  Current research projects are focussed on two areas (

  1. Determining how birds know what nest to build (behavioural and neurobiological laboratory work on zebra finches, behavioural field work on UK blue tits and African weavers);
  2. Using free-living hummingbirds (Canadian Rocky Mountains, collaborator Prof. Andy Hurly, University of Lethbridge) as a model system to investigate cognition in the wild.

Relevant references:

Subject area(s): Animal behaviour; Animal cognition

Keywords: Nest building, Birds, Foraging

Projects in Evolutionary Quantitative Genetics

Supervisor: Dr Michael Morrissey

Research area (s): Projects in Evolutionary Quantitative Genetics

Research description:Evolutionary quantitative genetics provides a general framework for modelling how natural selection and genetic variation interact to generate adaptive evolutionary change.  Conducting a research project in this field will provide a student with a solid foundation in a key area evolutionary biology, as well as broadly useful analytical skills.

Several projects are available that are suitable to an Msc by research, including:

  1. Selection of morphology and phenology in heterogeneous environments: The student will conduct a field study to collection trait, fitness, and microenvironmental data in a local wild annual plant population, and conduct analyses of selection of those traits that account simultaneously for the effects of both traits and microenvironmental variation on fitness.
  2. The genetic basis of plasticity and evolutionary consequences of non-linear reaction norms:  The student will collect trait data from fruit flies raised across a range of diet treatments.  This will allow inference of the genetic basis of plasticity, and assessment of likely changes in the mean and variance of phenotype under responses to different selective regimes.

Interested candidates should contact Dr Michael Morrissey to discuss these or other potential projects in advance of preparing an application.

Relevant references:

Subject area(s): Evolutionary Quantitative Genetics

Keywords: Genetics, Environmental, Evolutionary; Genetics

Predation, behaviour, animal communication

Supervisor: Professor Graeme Ruxton

Research area(s): Predation, behaviour, animal communication

Research description: I am interested in how prey protect themselves from predators, particularly through their appearance (e.g. camouflage and mimicry) and through grouping together. I have a number of projects in this area that should appeal to a student wanting to stretch their understanding in behavioural ecology, experimental design, animal behaviour and zoology generally. I also have a strong interest in enhancing the practice of experimental design and statistical analysis across biology, and would welcome enquiries from students with a maths and/or statistics background interested in applying their skills within whole organism biology.

Relevant references:

Subject area(s): Behavioural Ecology, Animal Communication

Keywords: Animal behaviour, Psychology, Camouflage, Signalling

Does dementia occur in sea mammals?

Supervisor: Professor Frank Gunn-Moore

Research area(s): Does dementia occur in sea mammals?

Research description:

A new Masters position is available to explore the question does dementia occur in sea mammals? Alzheimer’s disease in humans is a well-known phenomenon, but whether this form of dementia occurs in other mammals has only been explored in relatively small numbers, and very few other mammals have been potentially shown to have biochemical markers that look similar to those found in human patients (Youssef et al., 2016 Veterinary Pathology 53(2) 327-348). Recent data involving work from Florida, St Andrews and Oxford has suggested that these biomarkers may also be found in some species of stranded dolphins (Gunn-Moore et al., 2018 Alzheimer’s & Dementia, 14(2), 195-204). Therefore in this exciting and novel proposal due to a new collaboration between Prof Frank Gunn-Moore (Biomedical Science Research Complex, University of St Andrews), Prof Ailsa Hall (Director of the Sea Mammal Research Unit, University of St Andrews), Sir Prof Simon Lovestone (University of Oxford) and Dr Mark Dagleish (Moredun Research Institute, Edinburgh), we seek a candidate to explore a unique data base of sea mammal brains for the presence of potential biochemical changes that are associated with Alzheimer’s disease and other dementia. For enquiries, please contact:

Closing Date 1st May 2018.

Relevant References:

Subject area(s): Dementia, sea mammals

Keywords: dementia, Alzheimer’s, sea mammals

Genetics and sexual isolation

Supervisor: Professor Michael Ritchie

Research area(s): Genetics and sexual isolation

Research description: Finding genes that influence speciation is a major aim of evolutionary biology. We have identified genes with the potential to influence sexual isolation, the coordinated behaviours involved in species-specific mating choice. Species of Drosophila allow examination of mutants (some home-made) in these genes, and precise manipulation of gene expression. We offer a range of projects examining the behavioural consequences of altering these genes. Do mutants have abnormal courtship behaviour such as male songs or female preferences, court the wrong species, or suffer in sperm competition? Such projects allow behavioural, physiological and genetic studies of gene action and speciation.

Relevant References:

Wilkinson, G. S., et al. 2015. The locus of sexual selection: moving sexual selection studies into the post-genomics era. Journal of Evolutionary Biology 28: 739-755.

Ritchie, M. G. & Butlin, R. K. 2014. The genetics of insect mating systems. Pages 59-77 in Shuker, D. M. & Simmons, L. W. (eds.) The Evolution of Insect Mating Systems. OUP, Oxford.

Neville, M. C., et al. 2014. Male-specific fruitless isoforms target neurodevelopmental genes to specify a sexually dimorphic nervous system. Current Biology 24: 229-241.

Subject area(s): Evolutionary & Genetics; Speciation, sexual isolation, behaviour


Behavioural and Evolutionary Ecology of Insects

Supervisor: Dr David Shuker

Research area(s): Behavioural and Evolutionary Ecology of Insects

Research description: We study the behavioural ecology of insect reproduction. Our current research focuses on sexual behaviour in five species of seed bug (Family Lygaeidae). In particular, we are interested in instances of when “good mating systems go bad”, including heterospecific mating encounters (or “reproductive interference”) and failed copulations (“mating failure”). Both heterospecific matings and mating failure should be strongly disfavoured by natural and sexual selection, and yet both are more common that we have realised. Your project will explore one or both of these phenomena in our bugs, with a mix of behavioural and ecological experiments, grounded in mating systems theory.

Relevant References:

(1) Burdfield-Steel, E.R. & Shuker, D.M. (2011) Reproductive interference. Current Biology, 21: R450-451.

(2) Burdfield-Steel, E.R. & Shuker, D.M. (2014) The evolutionary ecology of the Lygaeidae. Ecology & Evolution, 4: 2278-2301.

(3) Greenway, E.V., Dougherty, L.R. & Shuker, D.M. (2015) Mating failure. Current Biology, 25: R534-R536.

(4) Greenway, E.V. & Shuker, D.M. (2015) The repeatability of mating failure in a polyandrous insect. Journal of Evolutionary Biology, 28: 1578-1582.

(5) Shuker, D.M., Currie, N., Hoole, T. & Burdfield-Steel, E.R. (2015) The extent and costs of reproductive interference among four species of true bug. Population Ecology, 57: 321-331.

(6) Shuker, D.M. & Simmons, L.W. (eds) (2014) The Evolution of Insect Mating Systems, Oxford University Press.

Subject area(s): Animal Behaviour, Behavioural Ecology

Keywords: Ecology, Evolution, Sexual Selection, Sexual Conflict

Pesticides and the costs to beneficial insects

Supervisor: Dr David Shuker

Research area(s): Pesticides and the costs to beneficial insects

Research description: Pesticides have an important role to play in helping us feed human populations. However, pesticides also bring negative effects, in terms of both human health and the health of the ecosystems around us. It is becoming increasingly clear that pesticides disrupt non-target species, often in ways that are more subtle than just killing them, but that still bring negative ecological consequences. We study the sub-lethal effects of controversial neonicotinoid pesticides on an important class of beneficial insects, the parasitic wasps. Your project will explore how neonicotinoids disrupt important life-history and behavioural decisions, such as sex allocation and mating.

Relevant References:

(1) Cook, N., Green, J., Shuker, D.M. & Whitehorn, P.R. (2016) Exposure to the neonicotinoid imidacloprid disrupts sex allocation cue use during superparasitism in the parasitoid wasp Nasonia vitripennis. Ecological Entomology, 41: 693-697.

(2) Ellis, C., Park, K., Whitehorn, P.R., David, A. & Goulson, D. (2017) The neonicotinoid insecticide thiacloprid impacts upon bumblebee colony development under field conditions. Environmental Science and Technology, 51: 1727-1732.

(3) Whitehorn, P.R., Cook, N., Blackburn, C.V., Gill, S.M., Green, J. & Shuker, D.M. (2015) Sex allocation theory reveals a hidden cost of neonicotinoid exposure in a parasitoid wasp. Proceedings of the Royal Society, Series B, 282: 20150389.

(4) Whitehorn, P.R., O’Connor, S., Wackers, F.L. & Goulson, D. (2012) Neonicotinoid pesticide reduces bumble bee colony growth and queen production. Science, 336: 351-352.

Subject area(s): Behavioural Ecology, Biological Control

Keywords: Ecology, Neonicotinoids, Parasitic Wasps, Sex Allocation

Computational biology

Supervisor: Dr V Anne Smith

Research area(s): Computational biology

Research description: Masters projects are available in areas of machine learning applied to molecular, neural, and ecological systems. Our group concentrates on inference of network structure from observational data, but also explores optimisation, agent-based modelling, and evolutionary algorithms, in the context of analysing biological questions. An emphasis is placed on evolutionary perspectives. Projects could be ideal bridges for students with degrees either in mathematics/computer science subjects or in biology to move into the interdisciplinary area of computational biology. Please contact Dr V Anne Smith ( to discuss your interests and particular projects which may suit. See below references for exemplars of research.

Relevant References: W Verleyen, SP Langdon, D Faratian, DJ Harrison, VA Smith. 2015. Novel Monte Carlo approach quantifies data assemblage utility and reveals power of integrating molecular and clinical information for cancer prognosis. Scientific Reports 5:15563

I Milns, CM Beale & VA Smith. 2010. Revealing ecological networks using Bayesian network inference algorithms. Ecology 91:1892-1899

C Echtermeyer, TV Smulders & VA Smith. 2010. Causal pattern recovery from neural spike train data using the Snap Shot Score. Journal of Computational Neuroscience 29:231-252

DJ White & VA Smith. 2007. Testing measures of animal social association by computer simulation. Behaviour 144:1447-1468

Subject area(s): Biological Sciences, Computer Science

Keywords: computational biology, bioinformatics, systems biology, machine learning

Experimental Evolution in Microbial Systems

Supervisor: Dr V Anne Smith

Research area(s): Experimental Evolution in Microbial Systems

Research description: How repeatable is evolution? What happens to synthetically engineered gene circuits under adaptive pressures? How do microbial communities persist? Masters projects are available in experimental evolution in microbial systems, particularly yeast. The laboratory has access to a state-of-the-art Bioscreen C machine which propagates 200 microbial cultures simultaneously. Topics addressed can range from basic features of evolution and mechanisms underlying adaptation, to exploration of robustness and persistence of biodiversity in artificial microbiomes, to impact of evolutionary considerations on design of systems for synthetic biology. Please contact Dr V Anne Smith ( to discuss your interests and particular projects which may suit.

Relevant References:

Subject area(s): Biological Sciences, Evolution

Keywords: experimental evolution, yeast, microbiomes, synthetic biology

Computational neuroscience: live imaging data

Supervisor: Dr V Anne Smith

Co Supervisor: Dr Stefan Pulver, School of Psychology and Neuroscience, University of St Andrews

Research area(s): Computational neuroscience: live imaging data

Research description: Live imaging of neural activity provides a wealth of data on neural activity in living animals; however, current computational analyses lag behind technological development. Masters projects are available in collaboration between a computational biologist (Smith) and an experimental neuroscientist (Pulver), developing and applying computational methods for ‘mining’ of live imaging datasets. Projects can address various aspects of analysis, from automatic image processing to answering biological questions by inferring neural information flow. Students can work entirely computationally, or have the opportunity to gain skills in experimental neuroscience. Please contact Dr V Anne Smith ( to discuss your interests and potential projects.

Relevant References:

Pulver SR, Bayley TG,, Taylor AL, Berni J, Bate M, Hedwig BJ. 2015. Imaging fictive locomotor patterns in larval Drosophila J. Neurophysiol. 114:2564-77

Lemon WC, Pulver SR, Höckendorf B, McDole K, Branson K, Freeman J, Keller PJ. 2015. Whole-central nervous system functional imaging in larval Drosophila. Nat. Commun. 11:7924

Echtermeyer C, Smulders TV, Smith VA. 2010. Causal pattern recovery from neural spike train data using the Snap Shot Score. Journal of Computational Neuroscience 29:231-252

Smith VA, Yu J, Smulders TV, Hartemink AJ, Jarvis ED. 2006. Computational inference of neural information flow networks. PLoS Computational Biology 2:e161.

Subject area(s): Neuroscience, Computer Science

Keywords: machine learning, live imaging, image processing, neuroinformatics

Agent-based modelling for animal behaviour

Supervisor: Dr V Anne Smith

Co Supervisor: Dr Lauren Guillette

Research area(s): Agent-based modelling for animal behaviour

Research description: Masters projects are available applying agent-based modelling to elucidate rules underlying animal cognition and behaviour. You will be jointly supervised by a computational biologist (Smith) with a background in animal behaviour, agent-based modelling and evolutionary programming, and an experimental biologist/psychologist (Guillette) whose current research concentrates on social learning during nest-building and foraging in birds, with interests in cognition and neurological bases of learning. Depending on student interest, projects may include working with extant data to build and evolve models, or incorporate significant amounts of hands-on behaviour experiments. Please contact Dr V Anne Smith ( to discuss projects which may suit.

Relevant References:

DJ White & VA Smith. 2007. Testing measures of animal social association by computer simulation. Behaviour 144:1447-1468.

VA Smith. 2008. Evolving an agent-based model to probe behavioral rules in flocks of cowbirds. Proceedings of the Eleventh International Conference on Artificial Life MIT Press, Cambridge, MA, pp 561-568.

Guillette, L.M., Scott, A.C.Y. & Healy, S.D. 2016 Social learning in nest-building birds: the role of familiarity. Proceedings of the Royal Society B: Biological Sciences, 283, 20152685.

Guillette, L.M. & Healy, S.D. 2017. The roles of vocal and visual interactions in social learning zebra finches: A video playback experiment. Behavioural Processes, 139, 43-49. doi:

Subject area(s): Behavioural Biology, Computer Science

Keywords: complex systems, social learning, animal behaviour, animal cognition

Evolutionary biology, developmental biology, cell biology, evo-devo, regeneration biology, genomics

Supervisor: Dr Ildiko Somorjai

Research area(s): Evolutionary biology, developmental biology, cell biology, evo-devo, regeneration biology, genomics

Research description: Have you ever wondered why some animals regenerate well, and humans do not? Are you interested in how new genes are born, and what generates diversity in animal body forms? The Somorjai lab addresses these problems from evolutionary, developmental and cell biological perspectives. We predominantly use the marine invertebrate chordate “amphioxus” due to its genetic and anatomical similarly to simple vertebrates. We also work on flatworms, which have amazing regenerative powers and multipotent stem cells. The project will depend on the student’s interests and background, but could include gene expression analyses, embryology, immunohistochemistry, confocal microscopy, genomics, and phylogenetic analyses.

Relevant References:

Bertrand S, Escriva H. Evolutionary crossroads in developmental biology: amphioxus. Development. 2011 Nov;138(22):4819-30.

Somorjai IM, Somorjai RL, Garcia-Fernàndez J, Escrivà H. Vertebrate-like regeneration in the invertebrate chordate amphioxus. Proc Natl Acad Sci U S A. 2012 109(2):517-22.

Dailey, SC, Planas, RF, Espier, AR, Garcia-Fernandez, J & Somorjai, IML Asymmetric distribution of pl10 and bruno2, new members of a conserved core of early germline determinants in cephalochordates. Frontiers in Ecology and Evolution. 2016. 3, 156.

Subject area(s): Evolutionary Biology, Developmental Biology

Keywords: Regeneration, Development, Evo-devo, Amphioxus

DNA repair in archaea and humans

Supervisor: Professor Malcolm White and Dr Carlos Penedo

Research area(s): DNA repair in archaea and humans

Research description: DNA repair is essential for all forms of life. There are many overlapping DNA repair pathways that contribute towards the maintenance of genetic integrity. This project will be focussed on improving our understanding of the Nucleotide Excision Repair (NER) pathway in humans. It will involve training in biochemistry and molecular biology, with an emphasis on the use of cutting-edge technigues to study DNA:protein interactions.

Relevant References:

Subject area(s): Molecular Biology, Biophysics

Keywords: DNA Repair, cancer, helicase, nuclease

Entry requirements and selection process

An undergraduate Honours degree at 2:1 level or better in biological or environmental sciences. Students from backgrounds such as mathematics may be accepted under exceptional circumstances.

If you studied for your first degree outside of the UK, please see the international entry requirements

For non-native English speakers, please see the English language requirements

Applicants will be short-listed by the project supervisor, and subject to interview by the project supervisor and an additional member of the Biology Postgraduate Committee.


For details of postgraduate tuition fees relevant to our research degrees including the MSc(Res), please visit:

Progression and assessment

Students in the MSc(Res) will be assigned an Internal Examiner (IE) and PG Tutor by the School. There will be a progress review meeting at three months to monitor and evaluate student progression, convened by the IE, with the student and Tutor in attendance. This meeting will be guided by a brief supervisor report and will be based on oral examination with no requirement for a written submission by the student.

The degree requires submission and examination of a dissertation at the end of the one-year program. As per 2016-2017 Senate Regulations (page 9), this thesis will consist of up to 30,000 words. The thesis will be evaluated by the IE and an External Examiner appointed at time of submission. Evaluation will be based on the written submission; there is no requirement for a viva.

Skills Training

In addition to the project-specific training that you will receive during your degree, Msc(Res) students will also have access to a wide range of training in transferable skills through the award-winning University of St Andrews GradSkills program, run by our Professional Development Unit CAPOD.

Specific post-graduate programs run within the School of Biology may also offer additional training, for instance in statistical, bioinformatics or molecular techniques.


Students may apply for placement in advertised projects or contact potential supervisors directly. We strongly recommend that potential candidates make contact with a potential supervisor before applying.  See links on this page.

Biology has two dates for admission to this degree: September and January each year.

If you have decided that you would like to make a formal application to study for an MSc(Res) at St Andrews, please complete an application using the online system.

Note: If you are self-funded and interested in working with a supervisor who does not currently have a project listed, please contact that person directly: supervisors’ email addresses may be found using the links on this page.