Graham Stone

Contact details



Ashworth 2 409
Institute of Evolutionary Biology
Ashworth Labs
Charlotte Auerbach Road

Post code


1986    BA First Class Zoology, Oxford University

1989    Ph.D. Oxford University Behavioural and physiological thermoregulation in solitary bees

1993-1998    Departmental Lecturer in Animal Biology, Oxford University Tutorial Fellow in Zoology, Magdalen College

1998-2005    Lecturer, Institute of Evolutionary Biology, Edinburgh University

2011-present    Professor of Ecology, Edinburgh University

Undergraduate teaching

I teach in several courses introducing the diversity and evolution of animal body plans, animal-plant interactions, and marine biology. I have contributed to tropical ecology courses in Kenya, run by the Tropical Biology Association, and co-authored a university level textbook on environmental physiology (The Environmental Physiology of Animals, by Pat Willmer, Graham Stone and Ian Johnston, Blackwells Science 1st Edn. 2000, 2nd Edn. 2005).

Open to PhD supervision enquiries?


Current PhD students supervised

Nora Villamil-Buenrostro (started April 2015) CoNaCyT Mexican scholarship program. Ecological costs of myrmecophily and its evolutionary consequences in a mexican ant plant, Turnera velutina.

Koorosh McCormack (started September 2017): The Processes of Natural Enemy Recruitment During Two Recent Gallwasp Invasions: Patterns emerging in Plagiotrochus quercusilicis and Dryocosmus kuriphilus.

Yusef Samari (started September 2017: The gardener’s dilemma: how should we manage urban green spaces to best benefit plants, pollinators, and people?

Past PhD students supervised

Dr. Rachel Atkinson (1997-2000). BBSRC quota studentship at Oxford University. Rachel is now a Ecosystem Restoration Scientist with Biodeiversity International.

Dr. Nigel Raine (1998-2001). Private Trust support, held jointly with Oxford University. Nigel is now Rebanks Professor in Pollinator Conservation at Guelph University, Canada.

Dr. Antonis Rokas (1998-2001). NERC quota award, ICAPB, Edinburgh University. Antonis is now Cornelius Vanderbilt Profeesor in Biological Sciences at Vanderbilt University in Nashville, Tennessee, USA.

Dr. Matthew Prescott (1999-2004). Leverhulme scholarship, Oxford University. Matt is now a campaigner for energy efficient and environmentally friendly technologies.

Dr. Alex Hayward (2002-2005). NERC quota studentship. Alex is now a BBSRC David Phillips Fellow in Biosciences at the University of Exeter

Dr. Martim Melo (2004-2006) Portuguese scholarship.Martim is now a postdoctoral researcher at the Research Centre in Biodiversity and Genetic Resources at the University of Porto, Portugal. He is an acknowledged expert on African birds.

Dr. Barbara Mackinder (2004-2006). Based at the Royal Botanic Gardens, Edinburgh. Barbara continues her work on botanical taxonomy as a member of the Centre for Middle eastern Plants at the Royal Botanic Gardens, Edinburgh.

Dr. Katherine Baldock (2005-2007). NERC quota studentship. Katherine is now a NERC Knowledge Exchange Fellow in Biological Sciences at the University of Bristol.

Dr. Richard Challis (2005-2007). NERC quota studentship. Richard is a postdoctoral informatician working on butterfly genomics with Mark Blaxter at the University of Edinburgh.

Dr. Tracey Begg (2005-2007). NERC quota studentship. Tracey is Policy & Advice Officer on Marine energy and seaweed harvesting for Scottish Natural Heritage.

Dr. Juan Carlos Ruiz-Guajardo (2006-2008). CONACYT- funded Mexican student. JC has just moved on from a postdoctoral research position at the university of California, Davis to a teaching position at Goshen College, Indiana.

Dr. Konrad Lohse (2008-2010). NERC quota studentship. Konrad is now a NERC indepdendent research fellow in population genomics at Edinburgh University.

Dr. Kate Armstrong (2006-2010). Based at the Royal Botanic Gardens, Edinburgh. Kate continues her work in plant taxonomy at the Institute of Systematic Botany, New York Botanical Garden.

Dr. Frazer Sinclair (2007-2011). Joint NERC quota studentship with CEH Wallingford. Frazer has worked on building conservation infrastructure in Sierra Leone and Indonesia, and is currently working on bird conservation on the island of Principe.

Dr. Jack Hearn (2009-2013). NERC quota studentship. After two spells of postdoctoral research in genomics and transcriptomics at Edinburgh University, Jack is about to start a new position on mosquito genomics at Liverspool School of Tropical Medicine.

Dr. Eugenio Valderrama (2011-2015) Darwin Trust. Eugenio is now a lecturer at El Bosque University, Bogotá, Colombia.

Julja Ernst (started 2011). NERC CASE studentship with CEH Wallingford. After the award of her Masters degree, Julja has set sail to run a mobile café, pretty much anywhere!

Dr. Tom Godfrey (2011-2015) BBSRC CASE studentship with Green Estate, Sheffield. Tom is about to start a position as a statistician with the Scottish Government.

Dr. Lisa Cooper (2013-2017) NERC quota studentship. Lisa has just started a senior technical position with a biotech company in Edinburgh.

Dr. Maria-Fernanda Torres (2014-2018) Darwin Trust. MaFe is about to start a post-doctoral position in the Antonelli Lab at the Department of Biological and Environmental Sciences, University of Gothenburg.

Research summary

I am a community ecologist with a strong focus on insect-plant interactions, combining molecular and field approaches to study of natural communities. I try to understand the role of evolutionary history in community composition, and in explaining why some species interact while others don't.

My research group incorporates DNA barcoding of species-rich communities, genomic approaches in the study of population history, and transcriptomic approaches to the study of gall induction.

You can see my research "in a nutshell" here:

I have three main projects on the go. 1. Assembly of cynipid gallwasp communities on oaks and related Fagaceae. 2. The evolution of associations between herbivores and neotropical trees in the genus Inga.  3. Quantification of the floral resources available to pollinators in urban and arctic habitats.

1. Assembly of cynipid gallwasp communities on oaks and related Fagaceae.

I have a long term research interest in the structure of parasitoid communities associated with the galls induced on plants by cynipid wasps. Ongoing projects address the assembly history of these communities across the world, the impact of gall traits on natural enemy assemblages, and identification of candidate genes involved in gall induction and development. I am particularly interested in how selection might generate the wide observed diversity of gall shapes, sizes and colours. We also  use invading gallwasps - and particularly the Oriental Chestnut gall wasp Dryocosmus kuriphilus - as model systems in which to study the recruitment of natural enemies to invading hosts.

Community structure work involves long collaborations with many people, particularly Karsten Schönrogge (CEH Wallingford), György Csoka (Hungarian Department of Forest Protection), George Melika (Plant Health and Molecular Biology Laboratory, Budapest), James Nicholls (Australian National Insect Collections, Canberra), Chang-Ti Tang (Smithsonian Institution, Washington DC), Jose-Luis Nieves-Aldrey (National Science Museum, Madrid) and Juli Pujade-Villar (University of Barcelona).

Population genetic work involves close collaboration with Konrad Lohse and Jack Hearn (Edinburgh University), Lynsey Bunnefeld (Stirling University), and Mike Hickerson (City College of New York).

Gall induction work has been led by Jack Hearn (Edinburgh University), and involves collaborations with Mark Blaxter (Edinburgh University), Jack Schultz (University of Toledo, USA), Joe Shorthouse (Laurentian University, Ontario) and Fredrik Ronquist (Swedish Museum of Natural History).

2. The evolution of associations between herbivores and neotropical trees in the genus Inga. 

This large collaborative project explores the impact of tree defensive chemistry and geography on herbivore associations, and builds on extensive work on Inga by the project leaders -  Tom Kursar and Lissy Coley  at the University of Utah, Salt Lake City. The project also involves Toby Pennington (Exeter University), Kyle Dexter (Edinburgh University), Maria-Jose Endara (University of Utah, Salt Lake City) and James Nicholls (Australian National Insect Collections, Canberra). 

3. Quantification of the floral resources available to pollinators in urban and other habitats.

My work on the value of urban habitats for pollinators began in the Urban Pollinator project, one of 9 projects in the multi-funder UK Insect Pollinators Initiative. Urban Pollinators was led by Jane Memmott (Bristol University) and brought together research teams in Edinburgh, Leeds (Bill Kunin) and Reading (Simon Potts). The work addressed key issues, including the relative abundance and species richness of flower visiting insects in urban agricultaural and nature reserve habitats, the value to pollinators of different urban habitat types, and the value of planted wildflower meadows for urban pollinators. This research opened my eyes to the need to understand how much floral resource (primarily nectar and pollen) is made available at the habitat and landscape scale. I have since extended work on the quantification of floral rewards to a range of planted wildflower meadows seed mixes, and (with Riikka Kaartinen, University of Helsinki) to arctic meadows at Zackenberg in northeastern Greenland.

Current research interests

I have three main projects on the go. 1. Assembly of cynipid gallwasp communities on oaks and related Fagaceae. 2. The evolution of associations between herbivores and neotropical trees in the genus Inga. 3. Quantification of the floral resources available to pollinators in urban and arctic habitats.Assembly of cynipid gallwasp communities on oaks and related Fagaceae. For further information on these, see 'Summary of research interests'.

Research activities

View all 13 activities on Research Explorer

Project activity

I have three main projects on the go, all with more information under the Research tab. 1. Assembly of cynipid gallwasp communities on oaks and related Fagaceae. 2. The evolution of associations between herbivores and neotropical trees in the genus Inga.  3. Quantification of the floral resources available to pollinators in urban and arctic habitats.

Current project grants

£40,000. 10/2017-4/2019. Lead PI. Fera-funded contract for research on the invading chestnut gallwasp, Dryocosmus kuriphilus, in Britain.

Past project grants

£250,000. 2/2015-4/2016. lead PI. Solicited one-off funding from BBSRC/Wellcome/NERC/DEFRA. Development of the Insect Pollinators Initiative specimen archive as a legacy for future research. Co-I’s Alfried Vogler (NHM London) and Adam Vanbergen (CEH, Edinburgh).

£60,229. 3/2014-3/2015. Lead PI. NERC Impact Acceleration Award, ‘Development of an Edinburgh Wildflower Meadows Seed Mix’, including funding from partners Scotia Seeds and Edinburgh City Council.

£110,307. 1/2014-1/2015. Leverhulme Grant (Ref RPG-2013-168). The phylogenetic origins of antiviral RNAi in animals Lead PI Darren Obbard (IEB Edinburgh), Co-PI Dr. Amy Buck (IIIR Edinburgh University)

£574,275. 1/2013 – 12/2015. NERC standard grant. Lead PI. Genomic approaches to inference of population history and multispecies community assembly. (NE/J010499). Co-PI’s James Cook (Reading University) and Konrad Lohse (IEB, Edinburgh University).

£1.25 million. 10/2011 – 10/2015. US National Science Foundation Dimensions of Biodiversity grant (£330,000 to Edinburgh). Coexistence, herbivore host choice, and plant-herbivore evolution in the recently radiated and speciose neotropical tree genus, Inga. Lead by Phyllis Coley and Tom Kursar (University of Utah), Toby Pennington and Catherine Kidner (Royal Botanic Gardens, Edinburgh).

£1.2 million. 1/2011 to 9/2014. Wellcome-BBSRC Insect Pollinator Initiative. Lead PI. Urban Pollinators: their ecology and conservation (BB/I000305/1). Consortium grant led by Jane Memmott (University of Bristol), Simon Potts (Reading) and Bill Kunin (Leeds).

£8,000. 1-6/2010. NERC NBAF pump-priming award (NBAF375 2009-2010). Lead PI. Nextgen sequencing approaches to population genetics. Co-I Jack Hearn (Edinburgh University).

£129,000. 1/2010 to 12/2012. INTEGRA-BIO Spanish Government award. Co-PI. Integrated analysis of the biodiversity, molecular systematics and macroecology of mediterranean insects. Lead PI José-Luis Nieves-Aldrey (Natural History Museum, Madrid).

£467,529. 1/2010-12/2012. NERC EHFI. Co PI. Climate change and management of forest biodiversity: predicting the impacts of climate matching strategies on plant-herbivore-enemy interactions. (NE/H000038/1). Joint award with Dr. Karsten Schönrogge (CEH Wallingford).

£575,353. 12/2007-12/2010. NERC standard grant. Lead PI. Using multispecies evolutionary history to test hypotheses of community assembly. (NE/E014453/1). Co-PI’s Andrew Rambaut and James Nicholls (IEB, Edinburgh University).

£65,500. 2/2006 to 10/2006. NERC small grant. Lead PI. Community-wide analysis of horizontal symbiont transmission: oak gallwasps as a model system. (NE/D007178/1). Co-PI Francis Jiggins (IEB, Edinburgh University), PDRA Alexandre Aebi.

£414,000. 12/ 2004 to 12/2007. NERC standard grant. Lead PI. Host-associated population substructure in generalist parasitoids: Oak cynipid gall communities as a model system (NER/B/504414/1). Co-P.I.s Sean Nee (IEB, Edinburgh University) and Karsten Schönrogge (CEH Wallingford).

£31,500. 3/2004-3/2007. NERC small grant. Lead PI. Guild phylogeography of oak-feeding gallwasps (NER/B/S2003/00856) (Co-P.I. Dr. Sean Nee, Edinburgh University)

£230,000. 6/2001 – 6/2004. NERC standard grant. Lead PI. Phylogenetic analyses of community structure: oak gallwasps as a model system. (GR/12847). Co- P.I.s James Cook (Imperial College at Silwood Park) and Karsten Schönrogge (CEH Wallingford).

£126,000. 9/1998 – 8/2001. NERC Ecological Dynamics of Genes thematic program. Lead PI. Inferring demographic processes using microsatellites: oak gallwasps as a model system (GST032035).

£130,000. 9/1998 – 8/2001. NERC standard grant. Co-PI. Partitioning of shared pollinators, and acacia-ant pollinator interactions, in highly seasonal acacia communities.(GR9/03553). Lead PI Pat Willmer, St. Andrews University.

View all 185 publications on Research Explorer