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Award details

University of Exeter funding: NERC GW4+ DTP PhD studentship

The form and function of dynamic courtship displays in bowerbirds. Biosciences PhD Studentship (NERC GW4+ DTP funded) Ref: 4006

About the award


Lead Supervisor

Dr Laura Kelley University of Exeter, Centre for Ecology & Conservation

Additional Supervisors

Dr Martin How University of Bristol, School of Biological Sciences

Professor Martin Stevens University of Exeter, Centre for Ecology & Conservation

Location: Penryn Campus, University of Exeter, Penryn, Cornwall.

This project is one of a number that are in competition for funding from the NERC Great Western Four+ Doctoral Training Partnership (GW4+ DTP).  The GW4+ DTP consists of the Great Western Four alliance of the University of Bath, University of Bristol, Cardiff University and the University of Exeter plus five Research Organisation partners:  British Antarctic Survey, British Geological Survey, Centre for Ecology and Hydrology,  the Natural History Museum and Plymouth Marine Laboratory.  The partnership aims to provide a broad training in earth and environmental sciences, designed to train tomorrow’s leaders in earth and environmental science. For further details about the programme please see

For eligible successful applicants, the studentships comprises:

  • An stipend for 3.5 years (currently £15,285 p.a. for 2020-21) in line with UK Research and Innovation rates
  • Payment of university tuition fees;
  • A research budget of £11,000 for an international conference, lab, field and research expenses;
  • A training budget of £3,250 for specialist training courses and expenses.
  • Up to £750 for travel and accomodation for compulsory cohort events.

Project details

Project Background

Male bowerbirds are famous for their complex courtship displays that incorporate a bower decorated with coloured objects, vocalisations, and vigorous display movements. Male great bowerbirds (Ptilonorhynchus nuchalis) also use their bowers to create visual tricks that affect the viewing female’s perception of the decorations that he presents during display. Despite the huge variety of signals that males produce when courting females, little is known about how the male’s physical display movements interact with his bower display, whether males modulate their displays in response to social and environmental changes, and the impacts on female mate choice. This project will quantify the female perspective of male displays to address these novel questions about interactions among display components, perception, and female choice.

Male bower 1

The male great bowerbird’s spectacular bower (© Laura Kelley)

Male bower 2

A male great bowerbird spends time tending to his bower while he waits for a female to arrive (© BBC Natural History Unit)

Project Aims and Methods

This project will increase our understanding of movement-based courtship displays and their interaction with an extended phenotype by addressing the following questions:

1. How do females perceive male displays? When displaying to females, males pick up, shake and toss a variety of coloured objects of two distinct shapes, interspersed with presentations of their pink crest. By videoing the male’s display from the female’s point of view from within the bower and using markerless object tracking software, this component of the project will quantify how the motion of both the male and the objects he displays are perceived by the female, how males vary in their displays, and whether females prefer males that produce more vigorous displays. 

2. How do display movements interact with other components of display? These experiments will address how the male’s display movements interact with the visual background and light environment by manipulating them and assessing changes in male displays. We will also use data from bower measurements and male displays to determine whether there are trade-offs or positive associations in the quality of different display components.

3. Do males improve their displays over time? Immature males spend up to seven years learning how to build bowers, and changes in display quality over time are poorly understood in this species. By quantifying the changes in courtship displays of individually identifiable males over multiple years, we will gain insights into the development of courtship behaviour, and will quantify the effects that social and environmental factors have on display quality.  

The questions addressed in this project can be adjusted where possible to take into account the successful candidate’s specific research interests.

This project would be ideal for a biologist with strong interests in sensory ecology, cognition and behaviour.  The successful candidate should demonstrate a willingness to learn new computational methods, be able to work under challenging field conditions and be a proactive problem solver. Previous fieldwork experience and a bird ringing license would be useful.


The student will receive training in the design and implementation of field experiments, video recording and analysis techniques including quantifying movements using 3D markerless tracking, modelling avian perception of colour and motion, statistical training, manuscript writing and publishing, science communication and outreach. 

Background reading and references

Kelley, LA & Endler, JA (2012) Illusions promote mating success in great bowerbirds. Science, 335, 335-338.

Endler, JA, Gaburro, J, Kelley, LA (2014) Visual effects in great bowerbird sexual displays and their implications for signal design. Proceedings of the Royal Society B, 281, 20140235.

Byers, J, Hebets, E, & Podos, J (2010) Female mate choice based upon male motor performance. Animal Behaviour, 79, 771-778. 


NERC GW4+ DTP studentships are open to UK and Irish nationals who, if successful in their applications, will receive a full studentship including payment of university tuition fees at the home fees rate.

A limited number of full studentships are also available to international students which are defined as EU (excluding Irish nationals), EEA, Swiss and all other non-UK nationals.  For further details please see the NERC GW4+ website.

Those not meeting the nationality and residency requirements to be treated as a ‘home’ student may apply for a limited number of full studentships for international students. Although international students are usually charged a higher tuition fee rate than ‘home’ students, those international students offered a NERC GW4+ Doctoral Training Partnership full studentship starting in 2021 will only be charged the ‘home’ tuition fee rate (which will be covered by the studentship). International applicants need to be aware that you will have to cover the cost of your student visa, healthcare surcharge and other costs of moving to the UK to do a PhD. More information on this is available from the universities you are applying to (contact details are provided in the project description that you are interested in

The conditions for eligibility of home fees status are complex and you will need to seek advice if you have moved to or from the UK (or Republic of Ireland) within the past 3 years or have applied for settled status under the EU Settlement Scheme.


Entry requirements

Applicants should have obtained, or be about to obtain, a First or Upper Second Class UK Honours degree, or the equivalent qualifications gained outside the UK.   Applicants with a Lower Second Class degree will be considered if they also have Master’s degree.  Applicants with a minimum of Upper Second Class degree and significant relevant non-academic experience are encouraged to apply.

All applicants would need to meet our English language requirements by the start of the  project


How to apply

In the application process you will be asked to upload several documents.  Please note our preferred format is PDF, each file named with your surname and the name of the document, eg. “Smith – CV.pdf”, “Smith – Cover Letter.pdf”, “Smith – Transcript.pdf”.

  • CV
  • Letter of application outlining your academic interests, prior research experience and reasons for wishing to undertake the project.
  • Transcript(s) giving full details of subjects studied and grades/marks obtained.  This should be an interim transcript if you are still studying.
  • If you are not a national of a majority English-speaking country you will need to submit evidence of your current proficiency in English.

Reference information
You will be asked to name 2 referees as part of the application process, however we will not expect receipt of references until after the shortlisting stage. Your referees should not be from the prospective supervisory team.

If you are shortlisted for interview, please ensure that your two academic referees email their references to the, 7 days prior to the interview dates.  Please note that we will not be contacting referees to request references, you must arrange for them to be submitted to us by the deadline.

References should be submitted by your referees to us directly in the form of a letter. Referees must email their references to us from their institutional email accounts. We cannot accept references from personal/private email accounts, unless it is a scanned document on institutional headed paper and signed by the referee.

All application documents must be submitted in English. Certified translated copies of academic qualifications must also be provided.

The closing date for applications is Friday 8 January 2021 2359 GMT .  Interviews will be held between 8th and 19th February 2021.  For more information about the NERC GW4+ DPT please visit

If you have any general enquiries about the application process please email  Project-specific queries should be directed to the lead supervisor.

Data Sharing
During the application process, the University may need to make certain disclosures of your personal data to third parties to be able to administer your application, carry out interviews and select candidates.  These are not limited to, but may include disclosures to:

  • the selection panel and/or management board or equivalent of the relevant programme, which is likely to include staff from one or more other HEIs;
  • administrative staff at one or more other HEIs participating in the relevant programme.

Such disclosures will always be kept to the minimum amount of personal data required for the specific purpose. Your sensitive personal data (relating to disability and race/ethnicity) will not be disclosed without your explicit consent.


Application deadline:8th January 2021
Value:£15,285 per annum for 2020-21
Duration of award:per year
Contact: PGR Enquiries