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Ecology and evolution of the earliest terrestrial animals. PhD in Biosciences (NERC GW4 + DTP)

  • Full or part time
  • Application Deadline
    Monday, January 06, 2020
  • Competition Funded PhD Project (European/UK Students Only)
    Competition Funded PhD Project (European/UK Students Only)

Project Description

Lead Supervisor:
Dr. Xiaoya Ma, Department of Biosciences, Centre for Ecology & Conservation, College of Life and Environmental Sciences, University of Exeter

Additional Supervisors:
Dr. Paul Kenrick, Department of Earth Sciences, the Natural History Museum
Dr. Greg Edgecombe, Department of Earth Sciences, the Natural History Museum
Dr. Erik Postma, Department of Biosciences, Centre for Ecology & Conservation, College of Life and Environmental Sciences, University of Exeter

Location: University of Exeter, Penryn Campus, Penryn, Cornwall TR10 9FE

This project is one of a number that are in competition for funding from the NERC GW4+ Doctoral Training Partnership (GW4+ DTP). The GW4+ DTP consists of the GW4 Alliance of research-intensive universities: the University of Bath, University of Bristol, Cardiff University and the University of Exeter plus five unique and prestigious Research Organisation partners: British Antarctic Survey, British Geological Survey, Centre for Ecology & Hydrology, the Natural History Museum and Plymouth Marine Laboratory. The partnership aims to provide a broad training in the Earth, Environmental and Life sciences, designed to train tomorrow’s leaders in scientific research, business, technology and policy-making. For further details about the programme please see

For eligible successful applicants, the studentships comprises:

- An stipend for 3.5 years (currently £15,009 p.a. for 2019/20) 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.
- Travel and accommodation is covered for all compulsory DTP cohort events.
- No course fee for courses run by the DTP

We are currently advertising projects for a total of 10 studentships at the University of Exeter

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 and Methods


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.


Examine the huge thin section collection of Rhynie chert material at the Natural History Museum in London, the Hunterian Museum, the University of Aberdeen, the University of Oxford and the University of Münster

Process field collections of Rhynie chert at the Natural History Museum

Using new imaging and analytical techniques to reveal the detailed structures of the organisms in the Rhynie chert, such as confocal laser scanning microscopy and synchrotron light source

Carrying out 3D reconstruction of these ancient organisms and their community

Using statistical methods to 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 project designs.

Funding Notes

NERC GW4+ funded studentship available for September 2020 entry. For eligible students, the studentship will provide funding of fees and a stipend which is currently £15,009 per annum for 2019-20.


References / Background reading list

- 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.
- Garwood et al. 2019. An introduction to the Rhynie chert. Geological Magazine. 10.1017/S0016756819000670
- Strullu-Derrien et al. 2016. A New Chytridiomycete Fungus Intermixed with Crustacean Resting Eggs in a 407-Million-Year-Old Continetal 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. PNAS, 115(10): E2274-E2283.
- Strullu-Derrien et al. 2019. Testate amoebae in the 407-Million-year-old Rhynie chert. Current Biology, 29 (3): 461-467.
- Ma et al. 2012. Complex brain and optic lobes in an early Cambrian arthropod. Nature, 490 (7419): 258-261.

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