Role for developmental modularity and integration in evolution of cranial diversity


This project will reveal the role for changes in modularity and integration in evolution of avian cranial diversity.

The studentship starts on the Oct 1 2018 and is funded by NERC. The deadline for application is Monday 8 January 2018.

How to apply

Read the eligibility criteria and application guidance below, then send your application to

Application deadline: 8 January 2018

Research focus

The aims of the project are:

The main goal for this PhD project is to survey this remarkable diversity morphometrically and to reveal the role for changes in modularity and integration in evolution of avian cranial diversity by studying species undergoing adaptive radiation.


This highly collaborative and multi-disciplinary PhD project will be a part of the larger effort to understand the nature and mechanisms of skull and beak diversity in modern birds. To successfully complete this project, this PhD student will need to learn how to conduct various types of morphometric analyses (Natural History Museum) as well as master laboratory-based techniques at Imperial College (Silwood Park campus).

3D CT (computer tomography)

To gain new insights into the nature of their diversification, this PhD student will join an active effort to analyse a growing database of 3D CT (computer tomography) scans of avian and other skulls in Abzhanov and Goswami groups.

More specifically, this student will perform analyses using morphometric landmarks to compare cranial shapes of all three clades (Darwin’s finches, honeycreepers and vangas) within the same morphospace.

The first aim is to quantitatively and qualitatively measure the degree of modularity and reveal patterns of integration of cranial skeletons in these examples of ecologically well-studied adaptive radiations and their relatives and outgroups. We will learn whether novel patterns of modularity or different levels of integration between cranial skeletal elements could be uncovered.

Such findings will help to better understand the nature of adaptive radiations in general and provide a foundation for future investigations on the molecular mechanisms underlying diversification of these morphologically distinguished groups of birds.

For the second aim, PhD student will analyse the basic mechanisms controlling integration between different skull modules (e.g. beak, orbits, cranial vault, etc) in the developing avian skull using chicken and zebrafinch embryos. Retroviral vectors and other experimental reagents will be used to reveal the nature of the molecular interactions between cranial bones in the skulls of live embryos.  These data will provide a foundation for future studies on the exact genetic causes of changes in modularity and integration during avian evolution.


This student will be trained to

  • study integration and modularity of the skull bones (using extensive expertise from Goswami group),
  • plot and analyze morphological data according to a particular phylogeny,
  • collect and process avian embryos,
  • clone multiple developmental genes for molecular tests,
  • perform immunohistological and in situ hybridization analyses
  • to image and analyse developmental gene expression (using experience and protocols of Abzhanov group).

For this project, extensive training will be provided to perform experiments with developing chicken and zebrafinch embryos.


Adaptive radiation is the rapid evolution of morphologically and ecologically diverse species from a single ancestor.

Comprising 30 orders and over 10,000 species, birds are the most successful land vertebrates and much of their success can be attributed to their cranial adaptations, such as uniquely shaped skull and beak.

The most famous examples of adaptive radiation in birds are Galapagos (Darwin’s) Finches (Thraupidae), Hawaiian honeycreepers (Drepanidinae) and Madagascar vangas (Vangidae), which all evolved remarkable levels of cranial morphological variation in sets of very closely related species making them ideal to study evolutionary and developmental mechanisms underlying such adaptive morphological radiations.


To be eligible for a full award a student must have:

  • Settled status in the UK, meaning they have no restrictions on how long they can stay.
  • Been ‘ordinarily resident’ in the UK for 3 years prior to the start of the studentship.
    This means they must have been normally residing in the UK (apart from temporary or occasional absences).
  • Not been residing in the UK wholly or mainly for the purpose of full-time education (This does not apply to UK or EU nationals).

How to apply

Applications for the PhD are processed through the Natural History Museum.

Applicants should apply directly to the lead supervisor Dr Arkhat Abzhanov with a CV and cover letter.  

The deadline for applications in 8 January 2018.

Any questions?

If you have any questions about the project please contact

Main supervisor: Dr Arkhat Abzhanov 

Science and Solutions for a Changing Planet (SSCP) doctoral training partnership

This is a joint project between The Science and Solutions for a Changing Planet (SSCP) Doctoral Training Partnership at Imperial College London and The Natural History Museum.

Funded by 

Submit your application