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Ecology and Conservation

Prof Ben Raymond

Prof Ben Raymond

Professor of Ecology and Evolution

 +44 (0)1326 259085

 SERSF 1:24


Science and Engineering Research Support Facility (SERSF):, University of Exeter,  Penryn Campus, Penryn, Cornwall, TR10 9FE, UK


I am an experimental evolutionary ecologist who works with insects, plants, bacterial pathogens and symbionts. Much of my research focuses on virulence and resistance, particularly the evolution of virulence in parasites and symbionts, the evolution of resistance to the biological control agent Bacillus thruringiensis and, increasingly, the evolution of resistance to antibiotics. I have also worked on the basic ecology of biocontrol agents and beneficial bacteria, such as B. thuringiensis, rhizobacteria and insect baculoviruses. As well as testing and advancing fundamental ideas, I am particularly interested in applying theory to real world problems such as improving the efficacy and sustainability of biocontrol tools or managing the evolution of resistance. My work ranges from the laboratory to the field and includes a number of in vivo and in vitro experimental evolution systems that have been developed by my group.


1998 DPhil (University of York)
1994 MSc (University College of North Wales)
1992 BA (St John’s College, Oxford)


2021 Professor of Ecology and Evolution, University of Exeter

2016-2021 Associate Professor of Microbial Ecology and Entomology, University of Exeter
2013-2106 Senior Lecturer in Evolutionary Ecology, Imperial College, Silwood Park.
2009-2013 NERC Research Fellow, Royal Holloway University of London.
2007-2009 NERC Research Fellow, University of Oxford
2005-2007 Insect pathogen evolutionary ecology, University of Oxford
2001-2004 Evolution of resistance to insect pathogens, Imperial College
1998-2001 Insect baculovirus ecology, CEH Oxford

Research group links

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Research interests

Cooperation and the evolution of virulence. The idea that kin selection might maintain group beneficial traits in microbes has revolutionized how we see virulence in bacteria, particularly virulence that is dependent on the extracellular export of proteins. My group has been one of the first to explore the implications of this theory in naturalistic host pathogen interactions, rather than in highly controlled artificial media.  While social interactions seem to be more important for some virulence factors than others, these ideas have been invaluable for understanding investment in virulence in B. thuringiensis and in other biocontrol agents such as entomopathogenic nematodes. Ongoing research projects are investigating whether how can apply our understanding of social interactions to shape the evolution of biocontrol agents in the laboratory and have resulted in a recent patent application.

Evolution and ecology of symbionts. Many bacteria switch between pathogenic, commensal and mutualistic lifestyles very readily over evolutionary timescales.  I am interested in the evolutionary ecology of selection for virulence in symbionts. We are also investigating plant-symbiont ecology and evolutionary ecology in a number of systems, including model crop species as well as the Titan Arum, Amorphophallus titanum, we have been exploring how environmental factors shaping the make-up of symbiont communities and whether this understanding can better shape our use of beneficial bacteria.

Evolution of resistance to biopesticides and GM crops. Applied as organic microbial pesticide, or as Cry toxins in GM crops, B. thuringiensis provides an exceptionally environmentally safe form of pest management, with no harmful effects on non-target organisms. It is therefore a technology worth preserving. This chiefly requires managing the evolution of resistance in target pests. Previously, I have explored how the fitness costs associated with resistance to B. thuringiensis could be manipulated to reduce the rate of evolution of resistance, as well as the value of biopesticide combinations in combating resistance. Current collaborations have investigated the genetic mechanisms of resistance in various Lepidopteran pests, and how insect behavior in the field might explain the very rapid evolution of resistance seen in some species. Recent research (with Oxitec and the University of Oxford) is investigating how the release of self-limiting genetically modified insects might be used as a tool to slow the evolution of resistance in diamondback moth, Plutella xylostella

Evolution of resistance to antibiotics. Resistance management is widely practiced in insect pest control but has had only a few successes in important human pathogens (mostly for the treatment of TB and HIV). Solutions to the current crisis in antibiotic resistance require imaginative solutions and better data. I am interested in applying many of lessons learnt in pesticide resistance management to the context of antibiotic resistance and have been developing a novel model experimental system (using Enterobacter cloacae in Lepidoptera) to test various ideas, including how best to use bacteriophage to combat the evolution of resistance.  

Biology and ecology of Bacillus thuringiensis. This bacterium is the world’s biggest selling microbial pesticide and supplies the key active proteins (Cry toxins) that are expressed in genetically modified insect resistant crops.   Despite its applied importance, the fundamental biology and ecology of this bacterium has been largely neglected, although this has broad implications for the biosafety of this organism and understanding how it kills its hosts. I am interested in how selection has led to the specialized production of large quantities of virulence factors and how this pathogen reproduces in the field in the absence of substantial epidemics. Ongoing interests include exploring the evolutionary relationship between B. thuringiensis and its complement of plasmids as well as evolutionary genomics and the importance of good science for understanding the biological safety of microbes in plant husbandry.


  • NERC Standard grant (2022-).   Managing the competition: how do burying beetles and microbes sustainably coexist in competition over shared resources? NE/V012053/1 (coI with Nick Royle)
  • British Academy (2020-2022) Inclusive and international risk assessment: Building a framework for gene drive organisms through collaboration, £200,000.  (co-I with Sarah Hartley)
  • Bayer CropScience LP.  Artificial selection for improvement of spore-forming entomopathogens. £159,150. PI Dec 2019.
  • Andermatt UK.  Field and laboratory testing of biopesticides based on Bacillus thuringiensis kurstaki.  £7,000 June 2019.
  • Horizon 2020 International Training Award “INSECT DOCTORS” proposal 859850, call H2020-MSCA-ITN-2019, co-investigator, €4.2 million €606,345 for Exeter
  • British Academy Knowledge Frontiers 2018, Co-developing risk assessment across disciplines and borders: gene drive mosquito field trials in Uganda.  coI with Sarah Hartley, £49,585.  KF2/100179.
  • BB/S002928/1 BBSRC industrial partnership award– ‘Group selection as a novel tool to screen and improve biological pesticides’ £942,085 (PI, with Neil Crickmore, Alistair Darby and DowDuPont) (2019-2021).
  • AHDB Studentship award  “Selection and improvement of insect pathogenic fungi for the control of multi-resistant aphids”  £71,400, (Oct 2018-June 2022)
  • EU RDF collaborative research award  “Microclimate and biocontrol interventions for Botrytis cinerea and fruit set problems in Cornish viticulture”  £127,170  (March 2018).
  • BBSRC Agrifood Catalyst £20,326 “Prototype development of novel bio-pesticide clones derived by experimental evolution” (Nov 2017)
  • Eden Collaboration Fund “Exploring plant microbial mutualisms in the pollination ecology of the Titan Arum, Amorphophallus titanum” £5,093 (March 2017)
  • MRC Innovation award. (PI) “Rapid assessment of phage for combating antimicrobial resistance in Enterobacter cloacae using a novel insect model.” £177,000 (May 2016). 
  • Leverhulme Trust. (PI) “Directing the evolution of bacterial virulence to overcome resistance to biological pesticides”.  £233,233.  RPG-2014-252 (Jan 2015).
  • BBSRC LINK grant Co-I .BB/L00819X/1 “Agricultural pest insect control: combining genetics, resistance management and dynamics”. £340,000 (March 2014)
  • Charles Sykes Trust £90720, co-I. “Antibiotic resistance in biofilms”(Sept 2011)
  • BBSRC Genome Analysis Centre, Capacity, Capability and Challenge Award, (PI) “ Identification of social and individually beneficial virulence genes in Bacillus thuringiensis via insertion site sequencing” £9996 (July 2011)
  • NERC Advanced Fellowship (NE/E012671/1) “Intra- and inter-specific competition and the evolution of cooperation in Bacillus thuringiensis”. £465,915 (Oct 2007)
  • BBSRC “Field and laboratory studies of Bacillus thuringiensis populations: the dynamics of diversity and its consequences for the evolution of resistance.” £303,269 (2005)  
  • BBSRC “The evolution of resistance to Bt toxins: testing ecological and evolutionary hypotheses in experimental microcosms.” £246,336 (2001)

Research networks

INSECT DOCTORS European Joint Doctoral Programme (EJD)

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Journal articles

Forsyth J, Barron N, Scott L, Bridget W, Chisnall M, Meaden S, Van Houte S, Raymond B (In Press). Decolonizing drug-resistant E. coli with phage and probiotics: breaking the frequency-dependent dominance of residents. Microbiology Abstract.
Raymond B, Lewis GC (In Press). Evaluation of endophyte infection of Lolium perenne on molluscan herbivory. Annals of Applied Biology (Supplement- Tests of Agrochemicals & Cultivars), 126, 104-105.
Morwool P, Dimitriu T, Crickmore N, Raymond B (In Press). Group selection as a basis for screening mutagenized libraries of public goods (Bacillus thuringiensis Cry toxins). Applied and Environmental Microbiology Abstract.
Raymond B, Matthews A, Erdos Z, Egleton M (In Press). Relative efficacy of biological control and cultural management for control of mollusc pests in cool climate vineyards. Biocontrol Science and Technology
White H, Sheppard S, Raymond B, Vos M, Pascoe B (In Press). Signatures of selection in core and accessory genomes indicate different. ecological drivers of diversification among Bacillus cereus clades. Molecular Ecology Abstract.
Erdos Z, Studholme DJ, Sharma MD, Chandler D, Bass C, Raymond B (2024). Manipulating multi-level selection in a fungal entomopathogen reveals social conflicts and a method for improving biocontrol traits. PLoS Pathog, 20(3). Abstract.  Author URL.
Penkova E, Raymond B (2024). When does antimicrobial resistance increase bacterial fitness? Effects of dosing, social interactions, and frequency dependence on the benefits of AmpC <i>β</i>-lactamases in broth, biofilms, and a gut infection model. Evolution Letters Abstract.
Williamson J, Matthews AC, Raymond B (2023). Competition and co-association, but not phosphorous availability, shape the benefits of phosphate-solubilizing root bacteria for maize (Zea mays). Access Microbiology, 5(12). Abstract.
Erdos Z, Studholme DJ, Raymond B, Sharma MD (2023). De novo genome assembly of Akanthomyces muscarius, a biocontrol agent of insect agricultural pests. Access Microbiology, 5(6). Abstract.
Dimitriu T, Souissi W, Morwool P, Darby A, Crickmore N, Raymond B (2023). Selecting for infectivity across metapopulations can increase virulence in the social microbe <i>Bacillus thuringiensis</i>. Evolutionary Applications, 16(3), 705-720. Abstract.
Raymond B, Erdos Z (2022). Passage and the evolution of virulence in invertebrate pathogens: Fundamental and applied perspectives. J Invertebr Pathol, 187 Abstract.  Author URL.
Erdos Z, Chandler D, Bass C, Raymond B (2021). Controlling insecticide resistant clones of the aphid, Myzus persicae, using the entomopathogenic fungus Akanthomyces muscarius: fitness cost of resistance under pathogen challenge. Pest Manag Sci, 77(11), 5286-5293. Abstract.  Author URL.
Matthews A, Majeed A, Barraclough TG, Raymond B (2021). Function is a better predictor of plant rhizosphere community membership than 16S phylogeny. Environ Microbiol, 23(10), 6089-6103. Abstract.  Author URL.
Hartley S, Smith RDJ, Kokotovich A, Opesen C, Habtewold T, Ledingham K, Raymond B, Rwabukwali CB (2021). Ugandan stakeholder hopes and concerns about gene drive mosquitoes for malaria control: new directions for gene drive risk governance. Malaria Journal, 20
Manktelow CJ, White H, Crickmore N, Raymond B (2020). Divergence in environmental adaptation between terrestrial clades of the Bacillus cereus group. FEMS Microbiol Ecol, 97(1). Abstract.  Author URL.
Erdos Z, Halswell P, Matthews A, Raymond B (2020). Laboratory sprayer for testing of microbial biocontrol agents: design and calibration. Abstract.
Zhou L, Slamti L, Lereclus D, Raymond B (2020). Optimal Response to Quorum-Sensing Signals Varies in Different Host Environments with Different Pathogen Group Size. mBio, 11(3). Abstract.
Manktelow CJ, Penkova E, Scott L, Matthews AC, Raymond B (2020). Strong Environment-Genotype Interactions Determine the Fitness Costs of Antibiotic Resistance. <i>In Vitro</i>. and in an Insect Model of Infection. Antimicrobial Agents and Chemotherapy, 64(10). Abstract.
Raymond BD, Zhou L, Somerville J (2019). Aseptic Rearing and Infection with Gut Bacteria Improve the Fitness of Transgenic Diamondback Moth, Plutella xylostella. Insects
Dimitriu T, Marchant L, Buckling A, Raymond B (2019). Bacteria from natural populations transfer plasmids mostly towards their kin. Proceedings of the Royal Society B: Biological Sciences, 286
Amanatidou E, Matthews AC, Kuhlicke U, Neu TR, McEvoy JP, Raymond B (2019). Biofilms facilitate cheating and social exploitation of β-lactam resistance in Escherichia coli. npj Biofilms and Microbiomes, 5(1). Abstract.
Amanatidou E, Matthews AC, Kuhlicke U, Neu TR, McEvoy JP, Raymond B (2019). Biofilms facilitate cheating and social exploitation of β-lactam resistance in Escherichia coli. NPJ Biofilms Microbiomes, 5(1). Abstract.  Author URL.
Patel M, Raymond B, Bonsall MB, West SA (2019). Crystal toxins and the volunteer's dilemma in bacteria. J Evol Biol, 32(4), 310-319. Abstract.  Author URL.
Raymond B (2019). Five rules for resistance management in the antibiotic apocalypse, a road map for integrated microbial management. Evolutionary Applications, 12(6), 1079-1091. Abstract.
Somerville J, Zhou L, Raymond B (2019). Gnotobiotic Rearing and Controlled Infection with Gut Symbionts Improve Adult Fitness in Transgenic Diamondback Moth, &lt;em&gt;Plutella xylostella&lt;/em&gt;. Abstract.
Ayra-Pardo C, Ochagavía ME, Raymond B, Gulzar A, Rodríguez-Cabrera L, Rodríguez de la Noval C, Morán Bertot I, Terauchi R, Yoshida K, Matsumura H, et al (2019). HT-SuperSAGE of the gut tissue of a Vip3Aa-resistant Heliothis virescens (Lepidoptera: Noctuidae) strain provides insights into the basis of resistance. Insect Sci, 26(3), 479-498. Abstract.  Author URL.
Dimitriu T, Medaney F, Amanatidou E, Forsyth J, Ellis RJ, Raymond B (2019). Negative frequency dependent selection on plasmid carriage and low fitness costs maintain extended spectrum β-lactamases in Escherichia coli. Scientific Reports, 9(1). Abstract.
Zhou L, Slamti L, Lereclus D, Raymond B (2019). Optimal response to quorum-sensing signals varies in different host environments with different pathogen group size. Abstract.
Matthews A, Pierce S, Hipperson H, Raymond B (2019). Rhizobacterial Community Assembly Patterns Vary Between Crop Species. FRONTIERS IN MICROBIOLOGY, 10  Author URL.
Matthews AC, Mikonranta L, Raymond B (2019). Shifts along the parasite-mutualist continuum are opposed by fundamental trade-offs. Proceedings of the Royal Society B: Biological Sciences, 286(1900). Abstract.
Mikonranta L, Buckling A, Jalasvuori M, Raymond B (2019). Targeting antibiotic resistant bacteria with phage reduces bacterial density in an insect host. Biol Lett, 15(3). Abstract.  Author URL.
Zhou L, Alphey N, Walker AS, Travers LM, Morrison NI, Bonsall MB, Raymond B (2019). The application of self-limiting transgenic insects in managing resistance in experimental metapopulations. Journal of Applied Ecology, 56(3), 688-698. Abstract.
Raymond B, Powell N (2019). We must follow these five rules to avert an antimicrobial resistance crisis. Pharmaceutical Journal, 303(7931).
Raymond B, Federici BA (2018). An appeal for a more evidence based approach to biopesticide safety in the EU. FEMS Microbiol Ecol, 94(1). Abstract.  Author URL.
Zhou L, Alphey N, Walker AS, Travers LM, Hasan F, Morrison NI, Bonsall MB, Raymond B (2018). Combining the high-dose/refuge strategy and self-limiting transgenic insects in resistance management-A test in experimental mesocosms. Evol Appl, 11(5), 727-738. Abstract.  Author URL.
Méric G, Mageiros L, Pascoe B, Woodcock DJ, Mourkas E, Lamble S, Bowden R, Jolley KA, Raymond B, Sheppard SK, et al (2018). Lineage-specific plasmid acquisition and the evolution of specialized pathogens in Bacillus thuringiensis and the Bacillus cereus group. Molecular Ecology, 27(7), 1524-1540. Abstract.
Dimitriu T, Marchant L, Buckling A, Raymond B (2018). Plasmid transfer is biased towards close kin in bacteria from natural populations. Abstract.
Geng LL, Shao GX, Raymond B, Wang ML, Sun XX, Shu CL, Zhang J (2018). Subterranean infestation by Holotrichia parallela larvae is associated with changes in the peanut (Arachis hypogaea L.) rhizosphere microbiome. Microbiological Research, 211, 13-20. Abstract.
Mikonranta L, Buckling A, Jalasvuori M, Raymond B (2018). Targeting antibiotic resistant bacteria with phages reduces bacterial density in an insect host. Abstract.
Zheng J, Gao Q, Liu L, Liu H, Wang Y, Peng D, Ruan L, Raymond B, Sun M (2017). Comparative Genomics of Bacillus thuringiensis Reveals a Path to Specialized Exploitation of Multiple Invertebrate Hosts. mBio, 8(4). Abstract.  Author URL.
Raymond B, Federici BA (2017). In defence of Bacillus thuringiensis, the safest and most successful microbial insecticide available to humanity—a response to EFSA. FEMS Microbiology Ecology, 93(7).
Medaney F, Ellis RJ, Raymond B (2016). Ecological and genetic determinants of plasmid distribution in Escherichia coli. Environ Microbiol, 18(11), 4230-4239. Abstract.  Author URL.
Shapiro-Ilan D, Raymond B (2016). Limiting opportunities for cheating stabilizes virulence in insect parasitic nematodes. Evolutionary Applications, 9(3), 462-470. Abstract.
Medaney F, Dimitriu T, Ellis RJ, Raymond B (2016). Live to cheat another day: Bacterial dormancy facilitates the social exploitation of β-lactamases. ISME Journal, 10(3), 778-787. Abstract.
Cornforth DM, Matthews A, Brown SP, Raymond B (2015). Bacterial Cooperation Causes Systematic Errors in Pathogen Risk Assessment due to the Failure of the Independent Action Hypothesis. PLoS Pathogens, 11(4). Abstract.
Deng C, Slamti L, Raymond B, Liu G, Lemy C, Gominet M, Yang J, Wang H, Peng Q, Zhang J, et al (2015). Division of labour and terminal differentiation in a novel Bacillus thuringiensis strain. ISME Journal, 9(2), 286-296. Abstract.
van Leeuwen E, O'Neill S, Matthews A, Raymond B (2015). Erratum: Making pathogens sociable: the emergence of high relatedness through limited host invasibility. The ISME Journal: Multidisciplinary Journal of Microbial Ecology, 9(10), 2328-2328.
Van Leeuwen E, O'Neill S, Matthews A, Raymond B (2015). Making pathogens sociable: the emergence of high relatedness through limited host invasibility. ISME Journal, 9(10), 2315-2323. Abstract.
Ayra-Pardo C, Raymond B, Gulzar A, Rodríguez-Cabrera L, Morán-Bertot I, Crickmore N, Wright DJ (2015). Novel genetic factors involved in resistance to Bacillus thuringiensis in Plutella xylostella. Insect Molecular Biology, 24(6), 589-600. Abstract.
Tellez-Rodriguez P, Raymond B, Moran-Bertot I, Rodriguez-Cabrera L, Wright DJ, Borroto CG, Ayra-Pardo C (2014). Strong oviposition preference for <i>Bt</i> over non-Bt maize in <i>Spodoptera frugiperda</i> and its implications for the evolution of resistance. BMC BIOLOGY, 12  Author URL.
Zhou L, Slamti L, Nielsen-LeRoux C, Lereclus D, Raymond B (2014). The social biology of quorum sensing in a naturalistic host pathogen system. Current Biology, 24(20), 2417-2422. Abstract.
Raymond B, Bonsall MB (2013). Cooperation and the evolutionary ecology of bacterial virulence: the Bacillus cereus group as a novel study system. BioEssays, 35(8), 706-716. Abstract.
Raymond B, Wright DJ, Crickmore N, Bonsall MB (2013). The impact of strain diversity and mixed infections on the evolution of resistance to Bacillus thuringiensis. Proceedings of the Royal Society B: Biological Sciences, 280(1769). Abstract.
Raymond B, West SA, Griffin AS, Bonsall MB (2012). The dynamics of cooperative bacterial virulence in the field. Science, 336(6090), 85-88. Abstract.
Garbutt J, Bonsall MB, Wright DJ, Raymond B (2011). Antagonistic competition moderates virulence in <i>Bacillus thuringiensis</i>. ECOLOGY LETTERS, 14(8), 765-772.  Author URL.
Raymond B, Wright DJ, Bonsall MB (2011). Effects of host plant and genetic background on the fitness costs of resistance to Bacillus thuringiensis. Heredity, 106(2), 281-288. Abstract.
Raymond B, Johnston PR, Nielsen-LeRoux C, Lereclus D, Crickmore N (2010). <i>Bacillus thuringiensis</i>: an impotent pathogen?. TRENDS IN MICROBIOLOGY, 18(5), 189-194.  Author URL.
Raymond B, Wyres KL, Sheppard SK, Ellis RJ, Bonsall MB (2010). Environmental factors determining the epidemiology and population genetic structure of the bacillus cereus group in the field. PLoS Pathogens, 6(5), 1-13. Abstract.
Martinou AF, Raymond B, Milonas PG, Wright DJ (2010). Impact of intraguild predation on parasitoid foraging behaviour. Ecological Entomology, 35(2), 183-189. Abstract.
Raymond B, Johnston PR, Wright DJ, Ellis RJ, Crickmore N, Bonsall MB (2009). A mid-gut microbiota is not required for the pathogenicity of <i>Bacillus thuringiensis</i> to diamondback moth larvae. ENVIRONMENTAL MICROBIOLOGY, 11(10), 2556-2563.  Author URL.
Raymond B, Ellis RJ, Bonsall MB (2009). Moderation of pathogen-induced mortality: the role of density in <i>Bacillus thuringiensis</i> virulence. BIOLOGY LETTERS, 5(2), 218-220.  Author URL.
Raymond B, Lijek RS, Griffiths RI, Bonsall MB (2008). Ecological consequences of ingestion of <i>Bacillus cereus</i> on <i>Bacillus thuringiensis</i> infections and on the gut flora of a lepidopteran host. JOURNAL OF INVERTEBRATE PATHOLOGY, 99(1), 103-111.  Author URL.
Bonsall MB, Raymond B (2008). Lethal pathogens, non-lethal synergists and the evolutionary ecology of resistance. JOURNAL OF THEORETICAL BIOLOGY, 254(2), 339-349.  Author URL.
Raymond B, Elliot SL, Ellis RJ (2008). Quantifying the reproduction of Bacillus thuringiensis HD1 in cadavers and live larvae of Plutella xylostella. Journal of Invertebrate Pathology, 98(3), 307-313. Abstract.
Raymond B, Davis D, Bonsall MB (2007). Competition and reproduction in mixed infections of pathogenic and non-pathogenic <i>Bacillus</i> spp. JOURNAL OF INVERTEBRATE PATHOLOGY, 96(2), 151-155.  Author URL.
Raymond B, Sayyed AH, Hails RS, Wright DJ (2007). Exploiting pathogens and their impact on fitness costs to manage the evolution of resistance to <i>Bacillus thuringiensis</i>. JOURNAL OF APPLIED ECOLOGY, 44(4), 768-780.  Author URL.
Raymond B, Sayyed AH, Wright DJ (2007). Host plant and population determine the fitness costs of resistance to Bacillus thuringiensis. Biol Lett, 3(1), 82-85. Abstract.  Author URL.
Raymond B, Hails RS (2007). Variation in plant resource quality and the transmission and fitness of the winter moth, <i>Operophtera brumata</i> nucleopolyhedrovirus. BIOLOGICAL CONTROL, 41(2), 237-245.  Author URL.
Raymond B, Sayyed AH, Wright DJ (2006). The compatibility of a nucleopolyhedrosis virus control with resistance management for Bacillus thuringiensis: Co-infection and cross-resistance studies with the diamondback moth, Plutella xylostella. Journal of Invertebrate Pathology, 93(2), 114-120. Abstract.
Raymond B, Sayyed AH, Wright DJ (2005). Genes and environment interact to determine the fitness costs of resistance to <i>Bacillus thuringiensis</i>. PROCEEDINGS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES, 272(1571), 1519-1524.  Author URL.
Raymond B, Hartley SE, Cory JS, Hails RS (2005). The role of food plant and pathogen-induced behaviour in the persistence of a nucleopolyhedrovirus. Journal of Invertebrate Pathology, 88(1), 49-57. Abstract.
Sayyed AH, Raymond B, Ibiza-Palacios MS, Escriche B, Wright DJ (2004). Genetic and biochemical characterization of field-evolved resistance to Bacillus thuringiensis toxin Cry1Ac in the diamondback moth, Plutella xylostella. Applied and Environmental Microbiology, 70(12), 7010-7017. Abstract.
Vanbergen AJ, Raymond B, Pearce ISK, Watt AD, Hails RS, Hartley SE (2003). Host shifting by <i>Operophtera brumata</i> into novel environments leads to population differentiation in life-history traits. ECOLOGICAL ENTOMOLOGY, 28(5), 604-612.  Author URL.
Raymond B, Vanbergen A, Watt A, Hartley SE, Cory JS, Hails RS (2002). Escape from pupal predation as a potential cause of outbreaks of the winter moth, Operophtera brumata. Oikos, 98(2), 219-228. Abstract.
Raymond B, Vanbergen A, Pearce I, Hartley SE, Cory JS, Hails RS (2002). Host plant species can influence the fitness of herbivore pathogens: the winter moth and its nucleopolyhedrovirus. Oecologia, 131(4), 533-541. Abstract.
Raymond B, Searle JB, Douglas AE (2001). On the processes shaping reproductive isolation in aphids of the Aphis fabae (Scop.) complex (Aphididae: Homoptera). Biological Journal of the Linnean Society, 74(2), 205-215. Abstract.
Raymond B, Darby AC, Douglas AE (2000). Intraguild predators and the spatial distribution of a parasitoid. OECOLOGIA, 124(3), 367-372.  Author URL.
Raymond B, Darby AC, Douglas AE (2000). The olfactory responses of coccinellids to aphids on plants. ENTOMOLOGIA EXPERIMENTALIS ET APPLICATA, 95(1), 113-117.  Author URL.
Raymond B (1999). Biological determinism unwarranted. Psycoloquy, 10 Abstract.


Raymond BD (2016). The biology, ecology and taxonomy of Bacillus thuringiensis and related bacteria. In Fiuza L, Polancyk RA, Crickmore N (Eds.) Bacillus thuringiensis and Lysinibacillus sphaericus: Characterization and use in the field of biocontrol, Springer. Abstract.
Raymond B, Wright DJ (2009). Resistance management of transgenic insect-Resistant crops: Ecological factors. In  (Ed) Environmental Impact of Genetically Modified Crops, 101-114. Abstract.

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External Engagement and Impact

Committee/panel activities

Member of DEFRA’s Advisory Committee on Releases to the Environment  (ACRE) which advises ministers on risks to the environments from releases of genetically modified organisms and biocontrol agents.

Member of Cornwall Antibioitic Resistance Group (CARG)

Invited lectures

Plenary speaker at: Siconbiol, Brazil 2015, SIP Vancouver  2015, International Organization for Biological Control, Innsbruck, 2011.  

Invited speaker: INTECOL symposium, London 2013; Bt evolution workshop Vienna 2012; European Spores meeting 2012; Biotechnology Congress, Cuba, 2011 & 2014. 

Invited talks: Liverpool, Edinburgh, Zurich, Jena, Roskilde, UNESP (Brazil), and Institutes of Plant Protection, Beijing & Wuhan.

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Module co-convenor for Zoology; co-convenor of Molecular Ecology module; co-ordinator of the MSci program.

Office hours: Tues 4-5pm & Friday 4-5pm



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Supervision / Group

Postdoctoral researchers

  • Michael Jardine postdoctoral fellow on NERC funded project exploring ecology and evolution of antimicrobial peptides in burying beetles

Postgraduate researchers

  • Hannah-Isadora Huditz
  • Loretta Mugo
  • Brogan Pett
  • Alessandro Roman
  • Hongbo Sun


  • Elli Amanatidou ; current employment Food Safety Authority
  • Tatiana Dimitriu : moved to postdoctoral positions in ESI now Royal Society research fellow
  • Zoltan Erdos Currently postdoctoral researcher in ESI with Angus Buckling
  • Jessica Forsyth Research Technician University of Manchester
  • James Manktelow ; Exeter graduate 2019
  • Andrew Matthews ; currently employed ESI University of Exeter
  • Frances Medaney ; moved to industry and post doctoral posts - currently employed by BBSRC (GCRF Hub)
  • Lauri Mikonranta : moved to post-doctoral position at the University of York with Ville Friman
  • Peter Morwool : currently working in industry in SW England in microbiological quality control
  • Elitsa Georgieva Penkova : currently PhD student in ESI with Anne Leonard
  • Hugh White : currently post doctoral researcher at Univeristy of Oxford
  • Liqin Zhou : PI, R

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Office Hours:

 Tues 4-5pm & Friday 3-4pm

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