Animal Behaviour and Physics PhD: How Do Bumblebee Colonies Regulate Nest Temperature Together?

Animal Behaviour and Physics PhD: How Do Bumblebee Colonies Regulate Nest Temperature Together?

Quick Decision

This Exeter Psychology PhD asks how a bumblebee colony’s collective behaviour and nest materials contribute to thermoregulation, and whether the nest can be understood as a biological metamaterial. The project explicitly allows empirical and modelling approaches to be developed and combined; it does not prescribe a sensor platform, colony count, experiment sequence, model family or software. Research would span the Exeter Bee Lab/CRAB and David Horsell’s CMRI laboratory, with Physics and Engineering co-supervisors. This is cross-department collaboration within a Psychology PhD, not an advertised joint degree. [S01, S04]

The team connects bumblebee behaviour with thermal and materials physics. A useful application should explain the applicant’s interest in linking biological questions to quantitative, experimental or modelling work. Before accepting a particular method as settled, ask what research gap the student will own, how biological and physical measurements will be combined, and what colonies, instruments and training are actually available. [S01, S08, S13]

The application deadline is noon on 8 December 2026. The project-specific award page gives 8 January 2027 for references, while the general scheme page gives 15 February; arrange references for the earlier date and ask PGR Admissions to confirm. The award lists an index-linked £21,805 stipend for 2026/27, up to 3.5 years of funding, tuition and £5,000 total research-training support. Confirm the indexed 2027/28 amount and international fee terms in writing. [S02, S03]

Research Problem and Doctoral Work

What is unresolved and why it matters

Social insect nests are not passive shelters. Individuals gather information from nestmates, nest structure and materials, and surrounding climate; their separate actions collectively shape shelter, internal conditions, maintenance and defense without central control. The project focuses on active bumblebee nests and their thermoregulation. It asks whether nest structures and materials function as biological metamaterials that manipulate heat, and how the colony maintains efficiency, equilibrium and resilience through social information, behavioral self-organization and use of space. The broader scientific idea is that material properties can emerge from movement and energy exchange among living components. [S01]

The timing is scientifically relevant. Field and laboratory studies show that colony heat regulation can incur energetic costs and may fail under high ambient temperatures, with consequences for brood, worker behavior and colony output. One 2024 heatwave experiment reported worker abandonment and fewer offspring when thermoregulation was unsuccessful; a 2023 study found that, in its experimental setup, colonies could regulate nest temperatures up to 30°C but not above that threshold despite increased energetic expenditure. A six-colony field study found that stronger colonies often held brood temperatures more steadily, while a 2026 study linked above-ground nesting to increased fanning demands under extreme projected summer heat. These results support studying colony-level feedbacks, but thresholds and outcomes differ by species, nest type and conditions. [S19, S20, S21, S23]

Work, methods and possible outputs

The advertised work is to investigate how bumblebee nest thermoregulation arises at the interface of physics, materials engineering and behavioral science. It explicitly names empirical and modelling approaches and says they may be developed and combined where appropriate. It also names social information, behavioral self-organization, spatial use, nest structure/materials and climatic conditions as relevant explanatory factors. The project aims to contribute to active matter and biometamaterials and to connect the findings to materials science, ecology and environmental monitoring. [S01]

The project PDF does not prescribe a sensor platform, species/colony count, controlled-temperature regime, data-analysis software, model family, sample size or fixed sequence of experiments. Recent field work shows that multi-sensor temperature monitoring and frequent sampling can reveal colony-specific patterns; heat-stress studies show why temperature, behavior and colony outcomes can be informative together. An analytically promising direction would be to link nest temperature or energy exchange with individual behavior and spatial allocation, then test whether structure or environmental variation alters stability. That is a research possibility suggested by the advertised aims and current literature, not a promised task or approved protocol. [S01, S19, S20, S21, S22, S23]

The novelty is credible but should be discussed precisely: the literature already studies bumblebee nest temperatures, heat stress, fanning, incubation and climate exposure. The opportunity is to connect those biological measurements with collective behavior, nest structure/material properties and active-matter or metamaterial concepts. A useful first conversation is therefore about the specific gap the team expects this student to own, and how the biological and physical measurements will be integrated. [S18-S23]

School, Programme and Field Context

The degree-awarding institution is the University of Exeter. The project PDF names a PhD in Psychology, Department of Psychology, at the Streatham campus. It places research in the Exeter Bee Lab within the Centre for Research in Animal Behaviour (CRAB), and says the student will spend time in the lab of second supervisor David Horsell at CMRI and interact with Physics and Engineering co-supervisors. This evidence supports cross-department collaboration; it does not establish a joint degree, double award or registration in CMRI's separate Physics/Engineering doctoral programme. [S01, S04, S10]

Exeter Psychology's research page describes four groups, including Animal Behaviour, a strong interdisciplinary approach and external partnerships. It reports that 92% of its REF 2021 research was rated 4* or 3*. In the 2021 national research assessment, Exeter's Psychology, Psychiatry and Neuroscience unit had 53% 4* and 39% 3* research; the Physics unit had 48% 4* and 45% 3*. The department page also reports 87th in THE Psychology subject rankings 2026 and 82nd in QS Psychology subject rankings 2026. These are useful subject-level signals, not a ranking of this project or evidence about supervision quality. [S05, S06, S07]

CRAB describes itself as a collaborative animal-behavior group with weekly group meetings and term-time seminars. CMRI provides a complementary materials-research environment and reports a large doctoral community, but its separately advertised PhD programme is in Physics/Engineering. For this project, the formal degree named in the PDF remains Psychology. Public materials do not specify whether the student joins CMRI's doctoral cohort, has access to every CMRI instrument, or follows a particular course/training schedule. [S08, S09, S10]

Supervisor and Lab Assessment

Professor Natalie Hempel de Ibarra is the lead supervisor and is listed as a CRAB academic and Psychology Director of Postgraduate Research. Her recent co-authored work provides useful evidence of the behavioral side of the project. Chapman et al. (2024) recorded bumblebee departures around sunrise and sunset and found that low-light conditions were associated with more hesitation or non-departure. Collett et al. (2023) analyzed how bumblebees coordinate body orientation and nest fixation in their first learning flight. Rands, Whitney and Hempel de Ibarra (2023) reviewed multimodal floral recognition, including temperature and other sensory cues. Together these papers show behavioral observation, orientation and sensory-learning expertise; they are adjacent to, rather than direct publications on, colony nest thermoregulation. [S08, S12, S15, S16, S17]

Dr David Horsell's work is in electrical and thermal conduction in novel materials, including graphene and nanostructures. A Royal Society case study describes an earlier interdisciplinary project on bumblebee hair as a thermal metamaterial, giving a relevant precedent for applying materials-science questions to bees. CMRI documents theory, design and modelling capabilities, including analytical and numerical tools and thermal-transport modelling. Those pages establish a relevant institutional skill base, but they do not say that a particular software package or instrument will be used in this studentship. [S09, S13, S14]

The named co-supervisors Steven Hepplestone and Matt Eames add Physics/Engineering links, but the public project description does not assign their specific roles. The main supervision questions are practical: who leads the biological and physical work; how often will the student meet each supervisor; what colonies, monitoring equipment and materials-characterization resources are available; and how much time is expected in CRAB versus CMRI? [S01, S08, S09, S14]

Funding, Application and Living Cost

Award and application conditions

The project sits in Exeter's competitive EPSRC Doctoral Landscape Award competition for September 2027. The award page lists up to 3.5 years full-time (part-time pro rata), an index-linked stipend currently £21,805 per year for 2026/27, University tuition fees, and a £5,000 Research Training Support Grant over the full 3.5 years—not £5,000 per year. The general scheme says up to 16 awards are available. The project-specific page states the university covers the home/international tuition-fee difference for successful international candidates; the general page says international awards are limited to up to 30% of the cohort. Visa, healthcare surcharge and moving costs are not covered. [S02, S03]

Submit by noon on Tuesday 8 December 2026 through the application route linked from the project description. The application requires an English personal statement, CV, full transcript(s), two references, and English proficiency evidence where applicable. Referees must email institutional letters. The date conflict matters: the project-specific Ref 6021 page says 8 January 2027, while the general EPSRC DLA page says 15 February 2027. Since an incomplete application may not progress, arrange references for 8 January and ask PGR Admissions which date controls. [S02, S03]

The scheme requires a first or upper-second UK honours degree, or equivalent, in an appropriate science/technology area; a lower-second may be considered with a Master's degree or significant relevant non-academic experience. The university also applies residence/fee-status rules, an international-award cap and English-language requirements where applicable. Because the project entry is 2027/28 but the current amount is explicitly a 2026/27 stipend rate, request the indexed 2027/28 figure and written confirmation of the award duration, stipend coverage and international fee waiver in the individual offer. [S02, S03]

Exeter living costs and stipend buying power

At the current published rate, £21,805 divided by 12 is about £1,817 per month. Exeter's April 2026 student budget estimate is £1,300–£2,000 per month, including accommodation, transport, food, utilities and personal/social costs. The simple comparison leaves roughly £517 per month at the low end and a shortfall of about £183 at the high end, before initial purchases or travel/relocation costs. The figure is a planning range, not a forecast of an individual budget, and the 2027/28 stipend and expenses may change. [S02, S24]

One current postgraduate room example is Cook Mews: £5,470.74 for a 44-week 2026/27 contract. The general budget page gives self-catered university halls at £150–£270 per week including bills; private rooms are £135–£250 per week without bills. Confirm what is included, whether a room is available for a 2027/28 doctoral student, the contract period, and the next-year price. International students should separately budget visa, healthcare surcharge, travel and start-up costs. [S03, S24, S25]

Applicant Fit and Preparation

Stated background and formal requirements

The project welcomes applicants from natural sciences, life sciences, physics or engineering, with an interest in modelling and/or experimental work applying physics-based solutions to biological or environmental problems. It states standard PhD entry requirements and does not name previous bee experience, a particular programming language, a specific laboratory technique or a particular modelling package as mandatory. Scheme-level academic, residency and English conditions still apply. [S01, S02, S03]

Evidence to prepare

Applicants can make their case more concrete by showing one example of quantitative research, experimental design, coding/modeling, animal behavior observation, thermal/materials measurement or work across disciplines. These are suggested ways to demonstrate readiness, not additional eligibility rules. A concise statement could explain (1) what can be measured at both colony and individual levels, (2) how empirical work could inform or test a model, and (3) how a Psychology registration can support a project that also spends time in Physics/Engineering facilities. The listed application asks for a personal statement, CV, transcripts and two references; document preparation should not wait on the conflicting reference date. [S01-S03]

In an interview, prepare to discuss the biological meaning of thermoregulation, how nests and social behavior could affect heat flow, and what evidence would distinguish a structural/material effect from a behavioral response. Be ready to ask how methods will be selected with the supervisors rather than pitching an over-specified design as if already agreed. The scheme indicates that shortlisted candidates may face a technical interview and successful nominees a general interview; the project page gives 15–24 February 2027 for the general interviews. [S02, S03]

Career Paths and Market Context

The most direct pathways are academic research in animal behaviour, behavioural ecology, collective systems or materials physics. Depending on the methods developed, transferable routes may also include environmental monitoring, scientific modelling and data analysis, or experimental and computational R&D in engineering and materials. The UK metamaterials roadmap describes applications across telecommunications, health, energy, transport, defence and manufacturing, while also identifying commercialisation, scale-up and skills challenges. Those are broad sector signals; they do not establish demand for work on bumblebee nest thermoregulation or this project’s graduate outcomes. The materials-engineer profile is an adjacent occupational reference, not a salary or placement forecast for this PhD. UK metamaterials roadmap · Materials engineer/scientist profile

Next Actions and Questions to Verify

Before applying

1. Which reference deadline is operative for Ref 6021: 8 January 2027 on the award page or 15 February 2027 on the general scheme page? Can PGR Admissions confirm by email, and should referees use a specific subject/reference format? [S02, S03]
2. What stipend amount is budgeted for September 2027 entry, and is the £5,000 total RTSG available across the full 3.5 years for this project? [S02, S03]
3. For applicants with international fee status, how many funded places are available in the competition, what residency rules apply to the applicant's circumstances, and does the award cover the full fee difference for the full award duration? [S02, S03]
4. Is an applicant from a particular adjacent discipline expected to demonstrate a particular prerequisite in experimental methods, physics, modelling or animal behavior beyond the stated standard PhD requirements? [S01, S02]

At interview

1. Which research question is the student's core responsibility: colony-level thermal performance, behavioral coordination/space use, nest structure/material properties, or integrating these scales? What is deliberately left open? [S01]
2. What methods are currently feasible and available: nest temperature/energy monitoring, behavioral tracking, materials characterization, controlled environmental treatments and/or computational modeling? Which are options rather than confirmed project tasks? [S01, S14, S19-S23]
3. How are responsibilities split across Professor Hempel de Ibarra, Dr Horsell, Dr Hepplestone and Dr Eames, and what is the expected supervision cadence and time in CRAB versus CMRI? [S01, S08-S14]
4. What training is planned for a Psychology PhD student crossing into materials/thermal physics, and which courses, facilities or research communities can the student actually access? [S01, S08-S10, S14]

Before accepting

1. Will the offer document explicitly confirm the degree registration as Psychology, award duration, stipend year/indexation, tuition-fee status, RTSG and any conditions for international students? [S01-S03, S10]
2. What 2027/28 accommodation costs and availability apply to the student's expected arrival date, and which utilities or one-off expenses remain outside rent? [S24, S25]
3. What career-development support is available for a doctoral researcher whose outputs may span animal behavior, ecological monitoring, physics-based modeling and materials research? Which outcomes are examples rather than placement commitments? [S10, S11, S26, S27]

Evidence and Sources

Project, programme and funding

• [S01] Current project description: source
• [S02] Ref 6021 award page: source
• [S03] General EPSRC DLA information: source
• [S04] Official 2027 project directory: source

University, subject and research environment

• [S05] Psychology research: source
• [S06] Psychology department and subject rankings: source
• [S07] Exeter REF 2021 results: source
• [S08] Centre for Research in Animal Behaviour: source
• [S09] Centre for Metamaterial Research and Innovation: source
• [S10] CMRI doctoral programme information: source
• [S11] CMRI alumni examples: source
• [S12] Psychology leadership: source
• [S13] Royal Society profile of David Horsell: source
• [S14] CMRI theory, design and modelling: source

Publications and field evidence

• [S15] Chapman et al. 2024: source
• [S16] Collett et al. 2023: source
• [S17] Rands, Whitney and Hempel de Ibarra 2023: source
• [S18] Ostwald et al. 2024: source
• [S19] Heatwave and colony-success study (2024): source
• [S20] Bretzlaff et al. 2023: source
• [S21] Gradišek et al. 2023: source
• [S22] Kevan, Rasmont and Martinet 2024: source
• [S23] Mullan et al. 2026: source

Costs and career context

• [S24] Exeter student budgeting: source
• [S25] Cook and Llewellyn Mews rates: source
• [S26] UK metamaterials commercialisation roadmap: source
• [S27] UK National Careers Service materials-engineer profile: source