Biological Sciences


Prof Graham Stone

Location: Ashworth 2 409

Telephone: 650 7194



Group members: No details available


1986BA First Class Zoology, Oxford University
1989Ph.D. Oxford University Behavioural and physiological thermoregulation in solitary bees
1993-1998Departmental Lecturer in Animal Biology, Oxford University Tutorial Fellow in Zoology, Magdalen College
1998-2005Lecturer, Institute of Evolutionary Biology, Edinburgh University
2011-presentProfessor of Ecology, Edinburgh University

Research groupings

Population Genetics Evolution, Animal Ecology and Behaviour


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), now in its second edition.

Research interests

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 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:

Research grants:

Currently funded research projects:

1.Genomic approaches to inference of population history and multispecies community assembly (NERC NE/J010499, 2013-2015) C with Dr Konrad Lohse(Edinburgh) and Prof James Cook (Reading and the University of Western Sydney).

2. Coexistence, herbivore host choice, and plant-herbivore evolution in the recently radiated and specioseneotropical tree genus, Inga (NSF, Dimensions in Biodiversity program, 2011-2014) C with Profs Phyllis Coley and Tom Kursar (University of Utah), Catherine Kidner(Edinburgh) and Prof Toby Pennington (Royal Botanic Gardens, Edinburgh)

3. Urban pollinators: their ecology and conservation (part of the multi-funder Insect Pollinators initiative, 2011-2014)- with Prof Jane Memmott (Bristol), Simon Potts (Reading), and Bill Kunin (Leeds).

Recently completed research projects:

1. Climate change and management of forest biodiversity: predicting the impacts of climate matching strategies on plant-herbivore-enemy interactions. (NERC NE/H000038/1, 2010-2012) - with Dr. Karsten Schonrogge (CEH Wallingford).

2. Using multispecies evolutionary history to test hypotheses of community assembly. (NERC NE/E014453/1, 2007-2010) - with Prof. A. Rambaut and Dr. J. Nicholls (Edinburgh).

3. Host-associated population substructure in generalist parasitoids: Oak cynipid gall communities as a model system. (NERC, NER/B/504406/1, 2004-2007) - with Dr Sean Nee (Edinburgh) and Dr. Karsten Schonrogge, CEH)

Community evolution and phylogeography

Oak insect communities: I use patterns of within-species genetic diversity to reconstruct the origins range expansion of species, using oak gallwasps and their chalcid parasitoids as a model system ( Stone et al. 2012, Lohse et al 2010, 2012, Nicholls et al. 2010, Stone et al 2007). A major current research question is whether the species making up these communities comprise sets of species with a long history of interaction, or recent assemblages from multiple origins. Our work involves using ABC and maximum likelihood analyses of multilocus genetic data, DNA barcoding ( Nicholls et al. 2012, Stone et al 2008) and study of bacterial endosymbionts such as Wolbachia and Rickettsia ( Weinert et al 2009, Rokas et al. 2002).

Neotropical Inga trees and their herbivores. Our NSF-fundedĀ  Dimensions in Biodiversity project focuses on the diversity and insect associates of the Neotropical tree genus Inga. Led by Profs. Lissy Coley and Tom Kursar at the University of Utah, the project explores the recent and rapid diversification of the tree genus Inga, and spatial patterns in their rich associated communities of herbivorous insects. The project combines transcriptome-driven phylogenetic analysis of the genus Inga, population genomic analysis of selected Inga species, and barcode-based analysis of herbivore communities in collaboration with Paul Hebert at > the Canadian Centre for DNA barcoding.

Pollination Ecology.

Urban Pollinators: My research group is part of the Urban Pollinators Project, a four-university consortium (with Leeds, Reading and Bristol) within the Inspect pollinators initiative that is exploring the distribution of pollinator biodiversity across the UK and exploring ways of enhancing pollinator diversity in Urban Habitats. I also have a PhD student, Tom Godfrey, working on related issues as a BBSRC CASE student with Green Estate, leading developers of planted pictorial meadows, based in Sheffield.>

Acacias and savannah pollination ecology. We have used plant-pollinator interaction networks to reveal the extent to which shared pollinators either facilitate pollination for multiple plant species, or mediate competition between them. This work, led by Dr. Kath Baldock, has focused in particular on savannah communities in Kenya, and the significance of daily pollinator activity patterns for network structure ( Baldock et al 2011). This research was based at Mpala Research Centre, Kenya and built on previous work on community pollination ecology in Tanzania ( Stone et al. 1998), Mexico (Raine et al. 2002, 2007) and Australia ( Gibson et al 2011). We have also used landscape genetic analysis in one acacia species ( Senegalia brevispica) to explore the spatial scale over which competition/facilitation for pollination may operate ( Ruiz-Guajardo et al 2010).

Phylogeny-based reconstruction of biological trait evolution

I am interested in the comparative method, and in its extension to analysis of patterns across populations within species ( Stone et al 2011). I have used multispecies phylogenies to test hypotheses about the evolution of defence against natural enemies ( Stone and Schonrogge 2003, Bailey et al 2009), animal reproductive strategies ( Stone et al. 2008), host plant associations ( Stone et al 2009, Cook et al. 2002), and bee thermal physiology ( Stone and Willmer 1989, Stone and Purvis 1992, Stone 1994).

Multitophicanalyses of oak insect community structure.

We use interaction webs to understand the structure of parasitoid-host and pollinator-plant interactions, and to examine the impacts of invading species on the structure of native communities. Recent work, based on the large INRA oak provenance trial at La Petite Charnie in France, has focused on the potential impacts of climate change on oak-herbivore-parasitoid interactions. Working in collaboration with Dr Karsten Schonrogge (Centre for Ecology and Hydrology), this NERC-funded research has sought to reveal the impact on trophic interactions of climate matched planting in forestry. This strategy seeks to plant seed crops (provenances) that are better matched than native trees to predicted future climates under predicted global change.

A long-running collaboration with Dr. Karsten Schonrogge has focused on multiple recent invasions of northwestern Europe by oak gallwasps (e.g. Schonrogge et al 2012, 2013) and the recent invasion of mediterranean Europe by the Chestnut gallwasp, Dryocosmus kuriphilus ( Gibbs et al 2011, Aebi et al 2007). We combine traditional identification techniques with DNA sequence 'barcodes', which allows us to link the eggs, larvae and adults of a single species and to separate biologically distinct but morphologically extremely similar species ( Acs et al 2010, Kaartinen et al 2010). I have a long running fascination with the biology of gallwasps in general, including ongoing discovery of many new species in Asia in collaboration with Chang-Ti Tang, a graduate researcher with Prof. Man-Miao Yang at the National Chung Hsing University, Taichung, Taiwan. (e.g. Tang et al 2011, Melika et al 2011).

Ant-plant interactions:

I have a long-term interest in ant-plant interactions ( Willmer et al 2009), and in particular in the potentially negative interactions between ant guards and pollinators. Though both are mutualists of a range of plants, the guards could potentially interfere with pollinator activity. We have shown that two ant-acacia mutualisms in America and Africa have evolved the same solution to the problem: short-lived floral ant repellents that keep ants away while the pollinators do their work ( Willmer and Stone 1997; Raine et al. 2002, 2004).

Environmental physiology: I use analyses of the thermal requirements of flight in bees and other insects to reveal the links between individual physiology and animal behaviour, particularly foraging and courtship.

Collaborations: I have much-valued long-standing collaborations with many highly respected taxonomists and field biologists, including Prof. Yoshihisa Abe (Kyushu University), Dick Askew(Emeritus), Gyorgy Csoka (Hungarian State Department of Forest Protection. Matrafured), George Melika (Budapest Plant Pest Diagnostic Laboratory), Jose-Luis Nieves-Aldrey (Museo Nacional deCiencias Naturales, Madrid), and Juli Pujade-Villar (Barcelona University).

Representative publications


Andrew RL, Bernatchez L, Bonin A, Buerkle CA, Carstens BC, Emerson BC, Garant D, Giraud T, Kane NC, Rogers SM, Slate J, Smith H, Sork VL, Stone GN, Vines TM, Waits L, Widmer A and Rieseberg LH (2013). A roadmap for molecular ecology. Molecular Ecology 22, 2605-2626.

Vanbergen AV et al. (2013) Threats to an ecosystem service: pressures on pollinators. Frontiers in Ecology and the Environment. Doi:

Pavlova, A., Amos, J. N., Joseph, L., Loynes, K., Austin, J. J., Keogh, J. S., Stone, G. N., Nicholls, J. A. and Sunnucks, P. (2013) Perched at the cyto-nuclear crossroads: divergent mitochondrial lineages correlate with environment in the face of ongoing nuclear gene flow in an Australian bird. Evolution (in press).

Schonrogge K, Begg T, and Stone GN (2013) Native birds and alien insects: spatial density dependence in songbird predation of invading oak gallwasps. PLOS One 5, 298-311.

Hickerson MJ, Stone GN, Lohse K, Demos T, Xia X and Takabayashi N (2013) Discrimination between a flawed method, and one that has merely been misapplied: a defense of msBayes and appropriate ABC model prior selection in response to Oaks et al. Evolution (in press).


Stone GN, Lohse K, Nicholls JA, Fuentes-Utrilla P, Sinclair F, Schonrogge K, Csoka G, Melika G, Nieves-Aldrey J-L, Pujade-Villar J, Tavakoli M, Askew RR and Hickerson MJ (2012). Reconstructing community assembly in time and space reveals enemy escape in a western palearctic insect community. Current Biology 22, 1-6.

Lohse K, Barton NH, Melika G and Stone GN (2012). A likelihood-based comparison of populations histories in a parasitoid guild. Molecular Ecology 21, 4605-4617.

Nicholls JA, Challis RJ, Mutun S and Stone GN (2012). Mitochondrial barcodes are diagnostic of shared refugia but not species in hybridising oak gallwasps. Molecular Ecology 21, 4051-4062.

Davies N, Field D and the Genomic Observatories Network (2012). A genomic network to monitor Earth. Nature 481, 145.

Schonrogge K, Begg T, Williams R, Melika G, Randle Z and Stone GN (2012). Range expansion and enemy recruitment by eight alien gallwasp species in Britain. Insect Conservation and Diversity. 5, 298-311.


Stone GN, Nee S and Felsenstein J (2011) Controlling for non-independence in comparative analysis of patterns across populations within species. Phil. Trans. Roy. Soc. Series B Biol Sci. 366,1410-1424.

Lohse K, Nicholls JA and Stone GN (2011) Inferring the colonisation of a mountain range - refugia vs. nunatak survival in high alpine ground beetles. Molecular Ecology 20, 394-408.60.

Gibbs M, Schonrogge K, Alma A, Melika G, Quacchia A, Stone GN and Aebi A (2011) Torymus sinensis: a viable management option for the biological control of Dryocosmus kuriphilus in Europe? BioControl 56, 527 C538.

Gibson M, Richardson DM, Marchante EM, Marchante HS, Rodger JG, Stone GN, Byrne M, Fuentes-Ramirez A, George N, Harris C, Johnson SD, Le Roux JJ, Miller, JT, Murphy D, Pauw A, Prescott MN, Wandrag E, and Wilson JRU (2011). Reproductive biology of Australian Acacia species: important mediator of invasiveness? Diversity and Distributions 17, 911-933.

Bihari P, Sipos B, Melika G, Feher B, Somogyi K, Stone GN and Penzes Z (2011) Western Palearctic phylogeography of an inquiline oak gallwasp, Synergus umbraculus. Biol J. Linn Soc. 102, 750 C764.

Lohse K, Sharanowski B, Blaxter M, Nicholls JA and Stone GN (2011) Developing EPIC markers for chalcidoid Hymenoptera from EST and genomic data. Molecular Ecology Resources 11, 521-529.

Baldock KCR, Memmott J, Roze D, Ruiz-Guajardo JC and Stone GN (2011) Daily temporal structure in African savannah flower-visitation networks, and consequences for network sampling. Ecology 92, 687 C698.

Melika, G., Tang, C.T., Nicholls, J., Yang, M.M. and Stone, G.N. (2011) Four new species of Dryocosmus gallwasps from Taiwan (Hymenoptera: Cynipidae, Cynipini). ISRN Zoology, doi:10.5402/2011/725180.

Tang, C-T, Melika G, Yang M-M, Nicholls J, and Stone GN (2011). A new genus of oak gallwasps, Cycloneuroterus Melika and Tang, with the description of five new species from Taiwan (Hymenoptera: Cynipidae: Cynipini). Zootaxa 3008, 33-62.

Tang, C-T, Melika G, Yang M-M, Nicholls J, and Stone GN (2011). New species of oak gallwasps from Taiwan (Hymenoptera: Cynipidae: Cynipini). Zootaxa 2865, 37 C52.


Lohse K, Sharanowski B and Stone GN (2010) Quantifying the Pleistocene history of the oak gall parasitoid Cecidostiba fungosa using twenty intron loci. Evolution 64, 2664 C2681.

Nicholls JA, Fuentes-Utrilla P, Hayward A, Melika G, Csoka G, Nieves-Aldrey J-L, Pujade-Villar J, Tavakoli M, Schonrogge K and Stone GN (2010) Community impacts of anthropogenic disturbance: natural enemies exploit multiple routes in pursuit of invading herbivore hosts. BMC Evolutionary Biology 2010, 10:322, 1-13.

Kaartinen R, Stone GN, Hearn J, Lohse K and Roslin T (2010). Revealing secret liaisons: DNA bar-coding changes our understanding of food webs. Ecological Entomology 35, 623-638.

Nicholls JA, Preuss S, Hayward A, Melika G, Csoka G, Nieves-Aldrey J-L., Askew RR, Tavakoli M, Schonrogge K and Stone GN (2010) Concordant phylogeography and cryptic speciation in two Western Palaearctic oak gall parasitoid species complexes. Molecular Ecology 19, 592 C609.

Acs, Z., Challis, R., Bihari, P., Blaxter, M., Hayward, A., Melika, G., Penzes, Z., Pujade- Villar, J., Nieves-Aldrey, J.-L., Schonrogge, K., and Stone, G.N. (2010). Phylogeny and DNA barcoding of inquiline oak gallwasps (Hymenoptera: Cynipidae) of the Western Palaearctic. Molecular Phylogenetics and Evolution 55, 210 C225.

Ruiz-Guajardo JC, Schnabel A, Ennos R, Preuss S, Otero-Arnaiz A and Stone GN (2010) Landscape genetics of the key African acacia species Senegalia mellifera (Vahl)- the importance of the Kenyan Rift Valley. Molecular Ecology 19, 5126-5139.

Melika, G., Pujade-Villar, J., Abe, Y., Tang, C.T., Nicholls, J., Wachi, N., Ide, T., Yang, M.M., Penzes, Zs., Csoka, Gy. and Stone, G.N. (2010) Palaearctic oak gallwasps galling oaks ( Quercus) in the section Cerris: re-appraisal of generic limits, with descriptions of new genera and species (Hymenoptera: Cynipidae: Cynipini). Zootaxa 2470, 1 C79.


Bailey R, Schonrogge K, Cook JM, Melika G, Csoka Gy, Thuroczy Cs, and Stone GN (2009). Host niches and defensive extended phenotypes structure parasitoid wasp communities. PLoS Biology, 7(8): e1000179.

Stone GN, Hernandez-Lopez A, Nicholls JA, di Pierro E, Pujade-Villar J, Melika G and Cook JM (2009).Extreme host plant conservatism during at least 20 million years of host plant pursuit by oak gallwasps. Evolution, 63, 854-869.

Penzes Z, Melika G, Bozsoki G, Bihari P, Miko I, Tavakoli M, Pujade-Villar P, Feher B, Fulandouml;p D, Szabo K, Bozso M, B Sipos, Somogyi K, and Stone GN (2009). Systematic re-appraisal of the gall-usurping wasp genus Synophrus Hartig, 1843 (Hymenoptera: Cynipidae: Synergini). Systematic Entomology, 34, 688-711.

Willmer PG, Nuttman CV, Raine NE, Stone GN, Pattrick JG, Henson K, Stillman, McIlroy L, Potts SG, and Knudsen JT (2009). Floral volatiles controlling ant behaviour. Functional Ecology, 23: 888-900.

Weinert LA, Werren JH, Aebi A, Stone GN and Jiggins FM (2009) Evolution and diversity of Rickettsia bacteria. BMC Biology 7, 6.

Tang, C-T, Melika G, Yang M-M, Nicholls J, Csoka Gy and Stone GN (2009). First record of an Andricus oak gallwasp from the Oriental Region: a new species from Taiwan (Hymenoptera: Cynipidae: Cynipini). Zootaxa 2175, 57 C65.


Stone GN, van der Ham RWJM, and Brewer JG (2008). Fossil oak galls preserve ancient multitrophic interactions. Proc. R. Soc. Lond. Ser. B. 275, 2213-2219.

Stone GN, Atkinson RJ, Rokas A, Nieves-Aldrey J-L, Melika G, Acs Z, Csoka G, Hayward A, Bailey R, Buckee C and McVean GAT (2008). Evidence for widespread cryptic sexual generations in apparently asexual Andricus gallwasps. Molecular Ecology 17, 652-665.

Tavakoli M, Melika G, Sadeghi SE, Penzes Z, Assareh MA, Atkinson RS, Bechtold M, Miko I, Zargaran MR, Aligolizade D, Barimani H, Bihari P, Fulop D, Somogyi K, Challis R, Preuss S, Nicholls J, and Stone GN (2008). New species of oak gallwasps from Iran (Hymenoptera: Cynipidae: Cynipini). Zootaxa, 1699, 1-64.

Garnier S, Verduijn M, Preuss S, Wolff K, Stone GN (2008). Polymorphic microsatellite loci and interspecific cross-amplification in the parasitoid wasps Megastigmus stigmatizans and Megastigmus dorsalis. Molecular Ecology Resources. 8, 421-424.


Stone GN, Challis RJ, Atkinson RJ, Csoka G, Hayward A, Mutun S, Preuss S, Rokas A, Sadeghi E, Schonrogge, K (2007). The phylogeographic clade trade: tracing the impact of human-mediated dispersal on the colonisation of northern Europe by the oak gallwasp Andricus kollari. Molecular Ecology 16, 2768-2781.

Challis RJ, Mutun S, Nieves-Aldrey J-L, Preuss S, Rokas A, Aebi A, Sadeghi E, Tavakoli M, and Stone GN (2007). Longitudinal range expansion and cryptic eastern species in the western Palaearctic oak gallwasp Andricus coriarius. Molecular Ecology 16, 2103-2114.

Acs Z, Melika G, Penzes Z, Pujade-Villar J and Stone, GN (2007). The phylogenetic relationships between Dryocosmus, Chilaspis and allied genera of oak gallwasps (Hymenoptera, Cynipidae: Cynipini). Systematic Entomology, 32, 70-80

van Noort S, Stone GN, Whitehead VB and Nieves-Aldrey, J.-L. (2007). Biology and redescription of Rhoophilus loewi (Cynipidae: Cynipoidea: Hymenoptera), with evolutionary implications on the inquilinism in gall wasps. Biological Journal of the Linnaean Society 90, 153-172.

van der Ham RWJM, Kuijper WJ Kortselius MJH, van der Burgh J, Stone GN, Brewer, JG (2008). Plant remains from the Kreftenheye Formation (Eemian) at Raalte, The Netherlands. Vegetation History and Archaeobotany 17, 127-144.

Raine NE, Sharp Pierson A, Stone GN (2007). Partitioning of pollinators during flowering in a Mexican Acacia community . A rthropod Plant Interactions 1, 101-117.

Ruiz-Guajardo JC, Otero-Arnaiz A, Taylor T, Stone GN, Glenn T, Schable NA, Miller JT and Schnabel A (2007). Molecular markers for the study of population genetics in Acacia mellifera and cross-species amplification of microsatellites within Acacia and closely related genera (Fabaceae: Mimosoideae). Molecular Ecology Notes 7, 1138-1140.

Aebi A, Schonrogge K, Melika G, Quacchia A, Alma A, Stone GN (2007). Native and introduced parasitoids attacking the invasive chestnut gall wasp Dryocosmus kuriphilus. EPPO Bulletin 37, 166-171.

Abe Y, Melika G and Stone GN (2007). The diversity and phylogeography of cynipid gallwasps (Hymenoptera, Cynipidae) of the Eastern Palaearctic and their associated communities. Oriental Insects, 41, 169-212.

Atkinson RJ, Rokas A and Stone GN (2007). Longitudinal patterns in species richness and genetic diversity in European oaks and oak gallwasps. Pp 127-154 in: Phylogeography in southern European refugia (ed. S. Weiss S). Springer, Dordrecht, The Netherlands.


Hayward A and Stone GN (2006). Comparative phylogeography across two trophic levels: the oak gall wasp Andricus kollari and its chalcid parasitoid Megastigmus stigmatizans. Molecular Ecology, 15, 479-489.

Kyriacou, H.M. Stone, G.N., Challis, R.J., Raza, A., Lyke, K.E., Thera, M.A., Kone, A.K., Doumbo, O.K., Plowe, C.V. and Rowe, J.A. (2006). Differential Var gene transcription in Plasmodium falciparum isolates from patients with cerebral malaria compared to hyperparasitaemia. Molecular and Biochemical Parasitology 150, 211-218.

Sadeghi SE,Melika G, Pujade-Villar J, Penzes Z, Acs Z, Bechtold M, Assareh MH, Tavakoli M, Yarmand H, Askary H, Stone GN, Azizkhani E, Zargaran MR, Aligolizade D, Barimani H and Dordaei AA (2006). Oak cynipid gall inquilines of Iran (Hym.: Cynipidae: Synergini), with descriptions of new species. Journal of the Entomological Society of Iran 25, 15-50.


Lowe A.J, Hicks BJ, Worley, K. Ennos RA, Morman JD, Stone, G and Watt AD (2005). Genetic differentiation in Scottish populations of the pine beauty moth, Panolis flammea. Bulletin of Entomological Research 95, 517-526.


Willmer PG and Stone GN (2004). Behavioral, ecological, and physiological determinants of the activity patterns of bees. Advances in the Study of Animal Behaviour 34, 347-466.

Raine NE, Gammans N, MacFadyen IJ, Scrivner GK and Stone GN (2004). Guards and thieves: antagonistic interactions between two ant species coexisting on the same ant plant. Ecological Entomology 29, 345-352.

Hale ML, Acs Z and Stone GN (2004). Polymorphic microsatellite loci in Eurytoma brunniventris, a generalist parasitoid in oak cynipid galls. Molecular Ecology Notes 4, 197-199.


Rokas A, Melika G, Abe Y, Nieves Aldrey J-L, Cook JM and Stone GN (2003). Lifecycle closure, lineage sorting and hybridisation revealed in a phylogenetic analysis of European oak gallwasps (Hymenoptera: Cynipidae: Cynipini) using mitochondrial sequence data. Molecular Phylogenetics and Evolution 26, 36-45.

Atkinson RJ, Brown GS and Stone GN (2003). Skewed sex ratios and multiple founding in galls of the oak apple gallwasp Biorhiza pallida (Hymenoptera: Cynipidae). Ecological Entomology 28, 14-24.

Rokas A, Atkinson, RJ, Webster L and Stone GN (2003). Out of Anatolia: longitudinal gradients in genetic diversity support a Turkish origin for a circum-Mediterranean gallwasp Andricus quercustozae. Molecular Ecology 12, 2153-2174.

Stone GN and French V (2003). Evolution: have wings come, gone, and come again? Current Biology 13, 436-438.

Stone GN and Schonrogge K (2003). The adaptive significance of insect gall morphology. Trends in Ecology and Evolution 18, 512-522.

Nee S and Stone GN (2003). The end of the beginning for neutral theory. Trends in Ecology and Evolution 18, 433-434.

Stone GN, Raine NE, Prescott P and Willmer PG (2003). The pollination ecology of Acacia. Australian Systematic Botany 16: 103-118.


Cook JM, Rokas A, Pagel M and Stone GN (2002). Evolutionary shifts between host oak sections and host plant organs in Andricus gallwasps. Evolution 56, 1821-1830.

Rokas A, Nylander JAA, Ronquist F and Stone GN (2002). A maximum likelihood analysis of eight phylogenetic markers using gallwasps (Hymenoptera: Cynipidae); implications for insect phylogenetic studies. Molecular Phylogenetics and Evolution 22, 206-219.

Raine NE, Willmer PG and Stone GN. (2002). Spatial structuring and floral repellence prevent ant-pollinator conflict in a Mexican ant-acacia Ecology 83, 3086 C3096.

Rokas A, Atkinson RJ, Nieves-Aldrey J-L, West, SA and Stone, GN (2002). The incidence and diversity of Wolbachia in gallwasps (Hymenoptera; Cynipidae) on oak. Molecular Ecology 11, 1815-1829.

Atkinson RJ, McVean GAT and Stone GN (2002). Use of population genetic data to infer oviposition behaviour: species-specific patterns in four oak gallwasps (Hymenoptera: Cynipidae). Proceedings of the Royal Society, Series B, 269, 383-390.

Stone GN, Schonrogge K, Atkinson RJ, Bellido D and Pujade-Villar J (2002). The population biology of oak gallwasps (Hymenoptera: Cynipidae). Annual Review of Entomology 47: 633-668.


Stone GN, Atkinson R, Rokas A, Csoka G, and Nieves-Aldrey J-L (2001). Differential success in northwards range expansion between ecotypes of the marble gallwasp Andricus kollari: a tale of two lifecycles . Molecular Ecology 10, 761-778.

Rokas A, Atkinson R, Brown G, West SA and Stone GN (2001). Understanding patterns of genetic diversity in the oak gallwasp Biorhiza pallida: demographic history or a Wolbachia selective sweep? Heredity 87, 294-305.


van Noort S and Stone GN (2000). Species richness, biogeography, conservation and pollination ecology of butterflies (Lepidoptera: Hesperioidea and Papilionoidea) in Mkomazi Game Reserve, Tanzania. Metamorphosis 11, 77-103.


Stone GN, Gilbert F, Willmer PG, Potts S, Semida F and Zalat S. (1999). Windows of opportunity and the temporal structuring of foraging activity in a desert solitary bee. Ecological Entomology 24, 208-221.


Stone GN, Willmer PG and Rowe JA (1998). Partitioning of pollinators during flowering in an African Acacia community. Ecology 79: 2808-2827.

Stone GN and Cook JM (1998). The structure of cynipid oak galls: patterns in the evolution of an extended phenotype. Proceedings of the Royal Society, Series B, 265, 979-988.


Willmer PG and Stone G (1997). Ant deterrence in Acacia flowers: how aggressive ant-guards assist seed-set. Nature 388, 165-167.

Willmer PG and Stone GN (1997). Temperature and water relations in desert bees. Journal of Thermal Biology 22, 453-465.


Schonrogge K, Stone GN and Crawley MJ (1996). Alien herbivores and native parasitoids: rapid development of guild structure in an invading gall wasp, Andricus quercuscalicis (Hymenoptera: Cynipidae). Ecological Entomology 21, 71-80.

Schonrogge K, Stone GN and Crawley MJ (1996). Abundance patterns and species richness of the parasitoids and inquilines of the alien gall former Andricus quercuscalicis Burgsdorf (Hymenoptera: Cynipidae). Oikos 77: 507-518.

Sunnucks PJ and Stone GN (1996). Genetic structure of invading insects and the case of the knopper gallwasp. Pp. 485-496 in: Frontiers of Population Ecology. CSIRO.

Stone GN, Willmer PG and Nee S (1996). Daily Partitioning of Pollinators in an Acacia Community. Proceedings of the Royal Society B, 263, 1389-1393.


Stone GN, Schonrogge K, Crawley MJ and Fraser S (1995). Geographic variation in the parasitoid community associated with an invading gallwasp, Andricus quercuscalicis (Hymenoptera: Cynipidae). Oecologia 104, 207-217.

Stone GN, Loder PMJ and Blackburn TM (1995). Foraging and courtship behaviour in males of the solitary bee Anthophora plumipes (Hymenoptera: Anthophoridae): thermal physiology and the role of body size. Ecological Entomology 20, 169-183.

Schonrogge K, Stone GN and Crawley MJ (1995). Spatial and temporal variation in guild structure: parasitoids and inquilines of Andricus quercuscalicis (Hymenoptera: Cynipidae) in its native and alien ranges. Oikos 72, 51-60.

Stone GN (1995). Female foraging responses to harassment in the solitary bee Anthophora plumipes. Animal Behaviour 50, 405-412.


Stone GN (1994b). Activity patterns of females of the solitary bee Anthophora plumipes in relation to temperature, nectar supplies and body size. Ecological Entomology 19, 177-189.

Stone GN (1994). Patterns of evolution of warm-up rates and body temperatures in flight in solitary bees of the genus Anthophora. Functional Ecology 8, 324-335.


Stone GN and Sunnucks PJ (1993). The population genetics of an invasion through a patchy environment: the cynipid gallwasp Andricus quercuscalicis. Molecular Ecology 2, 251-268.

Stone GN (1993a). Endothermy in the solitary bee Anthophora plumipes (Hymenoptera; Anthophoridae); independent measures of thermoregulatory ability, costs of warm-up and the role of body size. Journal of Experimental Biology 174, 299-320.

Stone GN (1993b). Thermoregulation in four species of tropical solitary bees: the roles of size, sex and altitude. Journal of Comparative Physiology B 163, 317-326.


Stone GN and Purvis A (1992). Warm-up rates during arousal from torpor in heterothermic mammals: physiological correlates and a comparison with heterothermic insects. Journal of Comparative Physiology B 162, 284-295.

Stone GN and Willmer PG (1989a). Endothermy and temperature regulation in bees: a critique of 'grab-and-stab' measurement of body temperature. Journal of Experimental biology 143, 211-223.

Stone GN and Willmer PG (1989b). Warm-up rates and body temperatures in bees: the importance of body size, thermal regime and phylogeny. Journal of Experimental Biology 147, 303-328.

Stone GN and Willmer PG (1989c). Pollination of Cardamom in Papua New Guinea. Journal of Apicultural Research 28, 228-237.

Willmer PG and Stone GN (1988). Pollination of lowland coffee ( Coffea canephora ); costs and benefits of leaf cutter bees in Papua New Guinea. Entomologia Experimentia et Applicata 50, 113-124.

Stone GN, Amos JN, Stone TF, Knight RJ, Gay H and Parrott F (1988). Thermal effects on activity patterns and behavioural switching in a concourse of foragers on Stachytarpheta mutabilis (Verbenaceae) in Papua New Guinea. Oecologia 77, 56-63.