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NERC E4 Roadside halophytes as a model for Anthropocene range shifts

   School of Biological Sciences

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  Dr R Milne, Dr A Twyford, Dr Markus Ruhsam  No more applications being accepted  Competition Funded PhD Project (Students Worldwide)

About the Project

Interested individuals must follow the "how to apply" link on the Geosciences E4 Doctoral Training Partnership web page:


Modern plants must respond to rapid anthropogenic habitat changes. The student will use molecular markers to examine two native coastal species that have recently moved inland along salted roadsides, asking how many genotypes were involved, and what adaptations were involved.

Project background

During the Anthropocene area, plants have been forced to adapt to rapid changes in their habitat, and future climate change coupled with accelerating human population expansion is likely to impose still more rapid and severe changes. One way to better understand how plants might adapt to this, or even be assisted to adapt, is to examine those plant species that have been able to expand their populations and range in response to the availability of new, anthropogenic habitats. However, large-scale human transport of alien species to new ranges obscures the signature from range expansion events that involve ONLY adaptation to new ranges. Therefore, the ideal study subjects are species that have expanded into new habitats that are adjacent to, and easily accessed from, their native range.

A key question here is, what challenges do plants face, to expand into novel habitats? Is it simply the case that a species must be pre-adapted to the new conditions, in which case any population could invade? Or might only certain genotypes have the required pre-adaptations? A third possibility is that at first no material is suitable to invade the new habitat, and that invasion only occurs when a suitable genotype arises. These hypotheses can be separated by examining both novel and original habitat populations using molecular markers, and determining how many separate invasion events there have been. 

Salting of roads has provided new habitats for salt-tolerant plant species previously confined to coastal habitats, that are easily accessible to those species. The British Isles provides an excellent study area within which to examine native plants that have invaded these two habitats, because of its large amounts of coastline and ample road coverage, plus a long tradition of botanical recording meaning that the timing of expansions is known with unusual precision. From this, we know that Cochlearia danica and Spergularia marina were largely confined to coastal regions before 1987, then rapidly expanded inland along roads. This project will examine the above questions using these two species as model systems, and then seek further understanding of these invasions at the genomic level.

Research questions

  1. Is all roadside material derived from native stock, or could some or all of it be alien?
  2. Assuming it is native, how many separate lineages of each species do the inland populations contain?
  3. Were there any genetic changes that occurred when the invasion process commenced, and if so did these enable invasion, or follow it?
  4. If the roadside invasion originated from certain coastal areas only, why was this the case?


Year 1: Literature review, starter collections and comparative analyses

  • Field collection of starter samples in autumn, then more extensive collection in spring and summer
  • Preliminary analyses of early collections, seeking and developing molecular markers capable of distinguishing between populations.
  • Review of relevant literature on biological invasions and range shifts.

Year 2: Fieldwork and genomics

  • Expansion of molecular marker survey to cover all sampled populations and refine markers to better separate material.
  • Genomic sequencing, seeking differences between inland and native populations at the genomic level.
  • Targeted fieldwork in spring and summer to expand sampling where there are gaps, or where molecular data indicates lineages of interest. Limited sampling outside Britain to provide outgroups and check for alien material.

Year 3: Integrated analysis of genomic data 

  • Detailed analysis of molecular data, modelling and mapping of range expansion process.
  • Analyses of genomic data to seek adaptive changes in inland material.
  • Writing up.


A comprehensive training programme will be provided comprising both specialist scientific training and generic transferable and professional skills. The project draws on the existing strengths of the University of Edinburgh, complemented by the Royal Botanic Garden Edinburgh, with its expertise in plant biodiversity science. Training will include: (1) analytical methods for genomic analysis, (2) data analysis, visualisation and programming in R, (3) plant study and fieldwork, (4) scientific writing and presentation, and (5) working in a multi-disciplinary environment. 


The student should be interested in using molecular population systematics to unravel the history of population expansion. Skills in, or a willingness to learn, bioinformatics will be important; an interest in biogeography and/or the UK flora is an advantage. There will be a lot of field collecting so field skills and a driving license would be advantages.

The School of Biological Sciences is committed to Equality & Diversity:

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Funding Notes

This project is eligible for the E4 Doctoral Training Partnership. The E4 projects are currently available for full NERC studentship funding which is competitive by interview to UK, EU and International applicants (The fee difference will be covered by the University of Edinburgh for successful international applicants.).
For application details see
Further details here -
Project Description –
Interested individuals must follow the "how to apply" link on the Geosciences E4 Doctoral Training Partnership web page:


The maps linked below are interactive. Click the legend and you can add/remove records from different time periods, to see the rapid spread:
BSBI (2022). BSBI Maps: Cochlearia danica., accessed 5th Oct 2022
BSBI (2022). BSBI Maps: Spergularia marina., accessed 5th Oct 2022

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