Ecology and evolution of the earliest terrestrial arthropods

This PhD project will investigate the invertebrate fauna preserved in the Early Devonian Rhynie chert.

The studentship is part of the Great Western Four+ Doctoral Training Partnership, funded by NERC and starts October 2020.

Apply for this course

Applications for this PhD are processed via the University of Exeter online application service.

Application deadline: 6 January 2020

Project background

The Rhynie chert is an Early Devonian geological site in Scotland (circa 407 million-years old), which preserves the oldest known terrestrial ecosystem, including plants, animals, fungi, algae and bacteria. The quality of fossil preservation in the Rhynie chert is astonishing, encompassing the most intact fossilised remains from a Palaeozoic terrestrial ecosystem, including preservation of cellular and ultrastructural details. Together, this fossil assemblage provides crucial insights into early life on land. 

In the past 100 years, research on the Rhynie chert has largely focused on its land plant-based components, but there are still major gaps in our understanding of its fauna and environment. In the past few years, the successful application of new imaging and analytical techniques (e.g. Confocal Laser Scanning Microscopy) have prompted fundamental breakthroughs in Rhynie chert fungal research, demonstrating the importance of interactions between fungi and the plant and animal elements of the system. 

Despite these major breakthroughs, we still have a poor understanding of what animals lived in the Rhynie chert communities and their roles in this early terrestrial ecosystem.

We will employ the latest imaging and analytical techniques to systematically describe and quantify the fauna of the Rhynie chert in order to gain an unprecedented picture of the biodiversity. In particular, we will investigate the nature of the interactions between different organisms and their palaeoenvironments, providing us with a unique reconstruction of this Palaeozoic terrestrial ecosystem. 

Project Aims

This research aims to determine the full biodiversity of the invertebrate fauna in the Rhynie chert, to reveal detailed anatomical structures of these animals and to identify their interactions with associated plants and fungi.

Consequently, these findings will contribute to our understanding of early land colonisation by invertebrates and will help to reconstruct the palaeoenvironment and palaeoecology of the Rhynie chert.

Project Methods

  • Examine the huge thin section collection of Rhynie chert material at the Natural History Museum in London, the Hunterian Museum, the University of Aberdeen and the University of Münster
  • Process field collections of Rhynie chert at the Natural History Museum
  • Using new imaging and analytical techniques to review the detailed structures of the organisms in the thin sections, such as confocal laser scanning microscopy
  • Carrying out 3D reconstruction of these ancient organisms and their community
  • Using statistical and other ecological research methods, investigate the palaeoecology of the Rhynie chert fossil bed

The doctoral researcher in this project is encouraged to assist in developing the overall research direction and experimental designs.


The researcher will receive training in palaeontologcial research methods, microscope-based imaging techniques and associated software (e.g., confocal laser scanning microscopy), statistical analysis, preparation of fossil materials and research and outreach work in museums. The student will have the opportunity to visit important museum and university collections in the UK and Germany.

All the skills they will learn are highly transferrable and will be useful to them irrespective of their precise career path within evolutionary biology and biological/environmental/palaeobiological sciences more generally.

Candidate Requirements

This project would suit a researcher with a passion for understanding evolutionary history. An excellent undergraduate degree in a relevant area is essential (e.g. biology, zoology, palaeontology), as is a desire for travel to view collections in the UK and Germany, an interest and ability to acquire skills in imaging, image analysis and statistical analysis, and excellent written and verbal communication skills.


Applications are primarily open to UK residents only (minimum residence of 3 years excluding time in further education), however, a limited number of full studentships are also available to EU residents. Students from EU countries who do not meet the residency requirements may still be eligible for a fees-only award but no stipend.  Applicants who are classed as International for tuition fee purposes are not eligible for funding.

All applicants need to comply with the registered university's English-language requirements.

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

How to apply

Applications for the PhD are processed via the University of Exeter online application service. You will need to include:

  • 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.
  • Two References (applicants are recommended to have a third academic referee, if the two academic referees are within the same department/school).

The deadline for applications is 6 January 2020.

Shortlisted candidates will then be invited to an institutional interview. Interviews will be held between 10 and 21 February 2020.

Visit the University of Exeter website for more information.

Any questions ?

You are encouraged to contact potential supervisors by email to discuss project-specific aspects of the proposed research at an early stage.

If you have any questions about the project please contact

Main supervisor: Dr Xiaoya Ma


University of Exeter

Main supervisor: Dr Xiaoya Ma

Co-supervisor: Dr Erik Postma

The Natural History Museum

Co-supervisor: Dr Greg Edgecombe

Co-supervisor: Dr Paul Kenrick


Edwards et al. 2017. History and contemporary significance of the Rhynie cherts—our earliest preserved terrestrial ecosystem. Philosophical Transactions of the Royal Society B, 373: 20160489.

Strullu-Derrien et al. 2016. A New Chytridiomycete Fungus Intermixed with Crustacean Resting Eggs in a 407-Million-Year-Old Continental Freshwater Environment. PLOS ONE 11: e0167301.

Dunlop & Garwood. 2017. Terrestrial invertebrates in the Rhynie chert ecosystem.  Philosophical Transactions of the Royal Society B, 373: 20160493.

Morris JL, et al 2018. The timescale of early land plant evolution. Proceedings of the National Academy of Sciences 115(10): E2274-E2283. (NE/N002067/1)

Ma et al. 2012. Complex brain and optic lobes in an early Cambrian arthropod. Nature, 490 (7419): 258-261.

Great Western Four+ Doctoral Training Partnership

Joint PhD training partnerships between the Natural History Museum and the Great Western Four, Bath, Bristol, Cardiff and Exeter universities.

Funded by