Liverpool Skylilne

Natural Sciences and Psychology

Dr Craig Wilding

Dr Craig Wilding

Telephone: 0151 231 2500


I have over twenty years of experience of the application of genetic and molecular biological techniques to the study of evolutionary biology, population genetics, human genetics and vector biology. I obtained a degree in Marine Biology from the University of Wales (Bangor) and subsequently completed my PhD research on the ‘Population genetics of the scallop Pecten maximus’. Following completion of my PhD I worked in evolutionary genetics at the University of Leeds before moving into Industry where I investigated human radiobiological genetics and genetic biomarkers of radiation exposure at Westlakes Research Institute. In 2006 I moved to the Liverpool School of Tropical Medicine (LSTM) to investigate the genetic mechanisms of insecticide resistance in mosquito vectors of human disease. After eight years at LSTM I joined LJMU as a lecturer in Evolutionary Genetics.

The main focus of my research is the genetic basis of insecticide resistance in mosquitoes. Resistance to the insecticides used in vector control represents a possible impediment to effective control strategies and an understanding of the genetic basis of this resistance would aid not only in the development of improved insecticide formulations, and hence more effective control measures, but also allow the development of genetic tests to rapidly measure the extent and spread of resistance. My work has used a variety of approaches including whole genome Agilent gene expression microarrays, qPCR, association mapping using the Illumina GoldenGate assay, targeted re-sequencing, and in vitro expression of identified candidates in bacterial and insect cell lines. I am particularly interested in methods to identify the regulatory variants underpinning the gene expression changes seen in insecticide resistant mosquitoes and to this end use reporter assays to study the activity of promoter sequences of candidate detoxification genes, and am involved in a project to characterise the microRNA repertoire (key post transcriptional regulators of genes) in An. gambiae (with Rodolphe Poupardin and Hilary Ranson, LSTM).

Recent work on insecticide resistance in An. gambiae and An. arabiensis has been conducted as part of the East African ICEMR (International Centre of Excellence for Malaria Research), based in Uganda and studying three key collection sites – Tororo, Jinja and Kanungu. I continue to have links with this programme. See

Additionally, I am involved, through the Donnelly group at LSTM, in a collaboration with the Kwiatkowski group at the Wellcome Trust Sanger Institute – the Anopheles 1000 genome initiative – undertaking whole genome sequencing of field-collected samples and lab colonies of Anopheles gambiae to study diversity and population structure of malaria vectors across a range of spatial scales in sub Saharan Africa. Through the Donnelly group I am also working on the annotation and analysis of insecticide resistance-associated gene families across a number of recently sequenced Anopheles mosquito species as part of the Anopheles genome cluster (see

I continue to have interests in the evolutionary genetics of reproductive isolation in the intertidal gastropod Littorina saxatilis. On some UK shores this snail exists as two morphotypes with a thin shelled morph (the H form) found high on the shore on cliffs and large boulders and a thick shelled morph (the M form) found low on the shore on and under boulders and on bedrock. Gene flow appears to be restricted, though possible, between these two forms. These snails are an interesting example of parapatric speciation and previous work utilising amplified fragment length polymorphism (AFLP) markers (Wilding et al. 2001) investigated the proportion of the genome responsible for this reproductive isolation. This was one of the first practical uses of outlier analysis to identify regions of the genome under selection. With the recent sequencing of the Littorina genome there is the potential to continue research on the genetic basis of this fascinating system.

Recently, I have also become interested in the genetics of reproductive isolation in two species of Tegenaria house spiders present throughout much of England and Wales. These species (Tegenaria saeva and T. gigantea) inhabit the west of England and Wales, and Eastern England respectively meeting in a hybrid zone which roughly follows the Welsh border.


1996, Bangor University, United Kingdom, Ph.D
1992, Bangor University, United Kingdom, BSc (Hons)


Journal Articles

Miles A, Harding NJ, Botta G, Clarkson CS, Antao T, Kozak K, Schrider DR, Kern AD, Redmond S, Sharakhov I, Pearson RD, Bergey C, Fontaine MC, Donnelly MJ, Lawniczak MKN, Kwiatkowski DP, Ayala D, Besensky NJ, Burt A, Caputo B, della Torre A, Fontaine MC, Godfrey HCJ, Hahn MW, Midega J, Neafsey DE, O'Loughlin S, Pinto J, Riehle MM, Vernick KD, Weetman D, Wilding CS, White BJ, Troco AD, Diabate A, Costantini C, Rohatgi KR, Besansky NJ, Elissa N, Coulibaly B, Dinis J, Midegal J, Mbogo C, Bejon P, Mawejje HD, Stalker J, Rockett K, Drury E, Mead D, Jeffreys A, Hubbard C, Rowlands K, Isaacs AT, Jyothi D, Malangone C, Vauterin P, Jeffery B, Wright I, Hart L, Kluczyriski K, Cornelius V, MacInnisn B, Henrichs C, Giacomantonio R. 2017. Genetic diversity of the African malaria vector Anopheles gambiae NATURE, 552 :96-+ >DOI >Link

Edmunds CV, Wilding CS, Rae R. 2017. Susceptibility of Chironomus plumosus larvae (Diptera: Chironomidae) to entomopathogenic nematodes (Rhabditida: Steinernematidae and Heterorhabditidae): Potential for control EUROPEAN JOURNAL OF ENTOMOLOGY, 114 :526-532 >DOI >Link

Silva Martins WF, Wilding CS, Steen K, Mawejje H, Antao TR, Donnelly MJ. 2017. Local selection in the presence of high levels of gene flow: Evidence of heterogeneous insecticide selection pressure across Ugandan Culex quinquefasciatus populations PLOS NEGLECTED TROPICAL DISEASES, 11 >DOI >Link

Martins WFS, Subramaniam K, Steen K, Mawejje H, Liloglou T, Donnelly MJ, Wilding CS. 2017. Detection and quantitation of copy number variation in the voltagegated sodium channel gene of the mosquito Culex quinquefasciatus SCIENTIFIC REPORTS, 7 >DOI >Link

Miles A, Harding NJ, Botta G, Clarkson C, Antao T, Kozak K, Schrider D, Kern A, Redmond S, Sharakhov I, Pearson R, Bergey C, Fontaine M, Troco A, Diabate A, Costantini C, Rohatgi K, Elissa N, Coulibaly B, Dinis J, Midega J, Mbogo C, Mawejje H, Stalker J, Rockett K, Drury E, Mead D, Jeffreys A, Hubbart C, Rowlands K, Isaacs A, Jyothi D, Malangone C, Vauterin P, Jeffrey B, Wright I, Hart L, Kluczynski K, Cornelius V, MacInnis B, Henrichs C, Giacomantonio R, Ayala D, Bejon P, Besansky N, Burt A, Caputo B, della Torre A, Godfray C, Hahn M, Neafsey D, O Loughlin S, Pinto J, Riehle M, Vernick K, Weetman D, Wilding C, White B, Lawniczak M, Donnelly M, Kwiatkowski D. 2016. Natural diversity of the malaria vector Anopheles gambiae bioRxiv, >DOI

Gomes B, Wilding CS, Weetman D, Sousa CA, Novo MT, Savage HM, Almeida APG, Pinto J, Donnelly MJ. 2015. Limited genomic divergence between intraspecific forms of Culex pipiens under different ecological pressures BMC EVOLUTIONARY BIOLOGY, 15 >DOI >Link

Weetman D, Mitchell SN, Wilding CS, Birks DP, Yawson AE, Essandoh J, Mawejje HD, Djogbenou LS, Steen K, Rippon EJ, Clarkson CS, Field SG, Rigden DJ, Donnelly MJ. 2015. Contemporary evolution of resistance at the major insecticide target site gene Ace-1 by mutation and copy number variation in the malaria mosquito Anophelesgambiae MOLECULAR ECOLOGY, 24 :2656-2672 >DOI >Link

Neafsey DE, Waterhouse RM, Abai MR, Aganezov SS, Alekseyev MA, Allen JE, Amon J, Arca B, Arensburger P, Artemov G, Assour LA, Basseri H, Berlin A, Birren BW, Blandin SA, Brockman AI, Burkot TR, Burt A, Chan CS, Chauve C, Chiu JC, Christensen M, Costantini C, Davidson VLM, Deligianni E, Dottorini T, Dritsou V, Gabriel SB, Guelbeogo WM, Hall AB, Han MV, Hlaing T, Hughes DST, Jenkins AM, Jiang X, Jungreis I, Kakani EG, Kamali M, Kemppainen P, Kennedy RC, Kirmitzoglou IK, Koekemoer LL, Laban N, Langridge N, Lawniczak MKN, Lirakis M, Lobo NF, Lowy E, MacCallum RM, Mao C, Maslen G, Mbogo C, McCarthy J, Michel K, Mitchell SN, Moore W, Murphy KA, Naumenko AN, Nolan T, Novoa EM, O'Loughlin S, Oringanje C, Oshaghi MA, Pakpour N, Papathanos PA, Peery AN, Povelones M, Prakash A, Price DP, Rajaraman A, Reimer LJ, Rinker DC, Rokas A, Russell TL, Sagnon N, Sharakhova MV, Shea T, Simao FA, Simard F, Slotman MA, Somboon P, Stegniy V, Struchiner CJ, Thomas GWC, Tojo M, Topalis P, Tubio JMC, Unger MF, Vontas J, Walton C, Wilding CS, Willis JH, Wu Y-C, Yan G, Zdobnov EM, Zhou X, Catteruccia F, Christophides GK, Collins FH, Cornman RS, Crisanti A, Donnelly MJ, Emrich SJ, Fontaine MC, Gelbart W, Hahn MW, Hansen IA, Howell PI, Kafatos FC, Kellis M, Lawson D, Louis C, Luckhart S, Muskavitch MAT, Ribeiro JM, Riehle MA, Sharakhov IV, Tu Z, Zwiebel LJ, Besansky NJ. 2015. Highly evolvable malaria vectors: The genomes of 16 Anopheles mosquitoes SCIENCE, 347 >DOI >Link

Wilding CS, Weetman D, Rippon EJ, Steen K, Mawejje HD, Barsukov I, Donnelly MJ. 2015. Parallel evolution or purifying selection, not introgression, explains similarity in the pyrethroid detoxification linked GSTE4 of Anopheles gambiae and An. arabiensis MOLECULAR GENETICS AND GENOMICS, 290 :201-215 >DOI >Link

Weetman D, Steen K, Rippon EJ, Mawejje HD, Donnelly MJ, Wilding CS. 2014. Contemporary gene flow between wild An. gambiae s.s. and An. arabiensis PARASITES & VECTORS, 7 >DOI >Link

Abdalla H, Wilding CS, Nardini L, Pignatelli P, Koekemoer LL, Ranson H, Coetzee M. 2014. Insecticide resistance in Anopheles arabiensis in Sudan: temporal trends and underlying mechanisms Parasites and Vectors, 7 >DOI

Chrismas NAM, Torres-Fabila B, Wilding CS, Grahame JW. 2014. An association of mitochondrial haplotype with shell shape in the intertidal gastropod Littorina saxatilis JOURNAL OF MOLLUSCAN STUDIES, 80 :184-189 >DOI >Link

Mitchell SN, Rigden DJ, Dowd AJ, Lu F, Wilding CS, Weetman D, Dadzie S, Jenkins AM, Regna K, Boko P, Djogbenou L, Muskavitch MAT, Ranson H, Paine MJI, Mayans O, Donnelly MJ. 2014. Metabolic and Target-Site Mechanisms Combine to Confer Strong DDT Resistance in Anopheles gambiae PLOS ONE, 9 >DOI >Link

Bass C, Zimmer CT, Riveron JM, Wilding CS, Wondji CS, Kaussmann M, Field LM, Williamson MS, Nauen R. 2013. Gene amplification and microsatellite polymorphism underlie a recent insect host shift Proceedings of the National Academy of Sciences, 110 :19460-19465 >DOI >Link

Mawejje HD, Wilding CS, Rippon EJ, Hughes A, Weetman D, Donnelly MJ. 2013. Insecticide resistance monitoring of field-collected anopheles gambiae s.l. populations from jinja, eastern uganda, identifies high levels of pyrethroid resistance Medical and Veterinary Entomology, 27 :276-283 >DOI

Mitchell SN, Stevenson BJ, Müller P, Wilding CS, Egyir-Yawson A, Field SG, Hemingway J, Paine MJI, Ranson H, Donnelly MJ. 2012. Identification and validation of a gene causing cross-resistance between insecticide classes in Anopheles gambiae from Ghana Proceedings of the National Academy of Sciences of the United States of America, 109 :6147-6152 >DOI

Weetman D, Wilding CS, Steen K, Pinto J, Donnelly MJ. 2012. Gene flow-dependent genomic divergence between Anopheles gambiae M and S forms Molecular Biology and Evolution, 29 :279-291 >DOI

Wilding CS, Smith I, Lynd A, Yawson AE, Weetman D, Paine MJI, Donnelly MJ. 2012. A cis-regulatory sequence driving metabolic insecticide resistance in mosquitoes: functional characterisation and signatures of selection Insect Biochemistry and Molecular Biology, 42 :699-707 >DOI

Jones CM, Liyanapathirana M, Agossa FR, Weetman D, Ranson H, Donnelly MJ, Wilding CS. 2012. Footprints of positive selection associated with a mutation (N1575Y) in the voltage-gated sodium channel of Anopheles gambiae Proceedings of the National Academy of Sciences of the United States of America, 109 :6614-6619 >DOI

Lima EP, Paiva MHS, De Arúajo AP, Da Silva EVG, Da Silva UM, De Oliveira LN, Santana AEG, Barbosa CN, De Paiva Neto CC, Goulart MO, Wilding CS, Ayres CFJ, De Melo Santos MAV. 2011. Insecticide resistance in Aedes aegypti populations from Ceará, Brazil Parasites and Vectors, 4 >DOI

Abilio AP, Kleinschmidt I, Rehman AM, Cuamba N, Ramdeen V, Mthembu DS, Coetzer S, Maharaj R, Wilding CS, Steven A, Coleman M, Hemingway J. 2011. The emergence of insecticide resistance in central Mozambique and potential threat to the successful indoor residual spraying malaria control programme Coleman M. Malaria Journal, 10 >DOI

Ayres CFJ, Müller P, Dyer N, Wilding CS, Rigden DJ, Donnelly MJ. 2011. Comparative genomics of the anopheline glutathione S-transferase epsilon cluster. PLoS One, 6 :e29237 >DOI >Link

Weetman D, Wilding CS, Steen K, Morgan JC, Simard F, Donnelly MJ. 2010. Association mapping of insecticide resistance in wild Anopheles gambiae populations: Major variants identified in a low-linkage disequilbrium genome PLoS ONE, 5 >DOI

Donnelly MJ, Corbel V, Weetman D, Wilding CS, Williamson MS, Black IV WC. 2009. Does kdr genotype predict insecticide-resistance phenotype in mosquitoes? Trends in Parasitology, 25 :213-219 >DOI

Wilding CS, Weetman D, Steen K, Donnelly MJ. 2009. High, clustered, nucleotide diversity in the genome of Anopheles gambiae revealed through pooled-template sequencing: Implications for high-throughput genotyping protocols BMC Genomics, 10 >DOI

Wilding CS, Weetman D, Steen K, Donnelly MJ. 2009. Accurate determination of DNA yield from individual mosquitoes for population genomic applications Insect Science, 16 :361-363 >DOI

Mzilahowa T, Ball AJ, Bass C, Morgan JC, Nyoni B, Steen K, Donnelly MJ, Wilding CS. 2008. Reduced susceptibility to DDT in field populations of Anopheles quadriannulatus and Anopheles arabiensis in Malawi: Evidence for larval selection Medical and Veterinary Entomology, 22 :258-263 >DOI

Wilding CS, Curwen GB, Tawn EJ, Sheng X, Winther JF, Chakraborty R, Boice JF. 2007. Influence of polymorphisms at loci encoding DNA repair proteins on cancer susceptibility and G(2) chromosomal radiosensitivity.(vol 48, pg 48, 2007) ENVIRONMENTAL AND MOLECULAR MUTAGENESIS, 48 :502-502 >DOI >Link

Bass C, Williamson MS, Wilding CS, Donnelly MJ, Field LM. 2007. Identification of the main malaria vectors in the Anopheles gambiae species complex using a TaqMan real-time PCR assay Malaria Journal, 6 >DOI

Wilding CS, Curwen GB, Tawn EJ, Sheng X, Winther JF, Chakraborty R, Boice JD. 2007. Influence of polymorphisms at loci encoding DNA repair proteins on cancer susceptibility and G2 chromosomal radiosensitivity. Environ Mol Mutagen, 48 :48-57 >DOI >Link


Wilding CS, Trikic MZ, Hingston JL, Copplestone D, Janet Tawn E. 2006. Mitochondrial DNA mutation frequencies in experimentally irradiated compost worms, Eisenia fetida Mutation Research - Genetic Toxicology and Environmental Mutagenesis, 603 :56-63 >DOI

Wilding CS, Cadwell K, Tawn EJ, Relton CL, Taylor GA, Chinnery PF, Turnbull DM. 2006. Mitochondrial DNA mutations in individuals occupationally exposed to ionizing radiation. Radiat Res, 165 :202-207 >Link

Wilding CS, Rees GS, Relton CL, Tawn EJ. 2006. Genotype profiles of loci encoding DNA repair enzymes in newborn and elderly populations: no evidence of association with longevity. Biogerontology, 7 :35-41 >DOI >Link

Grahame JW, Wilding CS, Butlin RK. 2006. Adaptation to a steep environmental gradient and an associated barrier to gene exchange in Littorina saxatilis. Evolution, 60 :268-278 >Link

De Ruyck K, Wilding CS, Van Eijkeren M, Morthier R, Tawn EJ, Thierens H. 2005. Microsatellite polymorphisms in DNA repair genes XRCC1, XRCC3 and XRCC5 in patients with gynecological tumors: Association with late clinical radiosensitivity and cancer incidence Radiation Research, 164 :237-244 >DOI

Wilding CS, Relton CL, Rees GS, Tarone RE, Whitehouse CA, Tawn EJ. 2005. DNA repair gene polymorphisms in relation to chromosome aberration frequencies in retired radiation workers. Mutat Res, 570 :137-145 >DOI >Link

Simpson RJ, Wilding CS, Grahame J. 2005. Intron analyses reveal multiple calmodulin copies in Littorina. J Mol Evol, 60 :505-512 >DOI >Link

Relton CL, Wilding CS, Pearce MS, Laffling AJ, Jonas PA, Lynch SA, Tawn EJ, Burn J. 2004. Gene-qene interaction in folate-related genes and risk of neural tube defects in a UK population Journal of Medical Genetics, 41 :256-260

Relton CL, Wilding CS, Laffling AJ, Jonas PA, Burgess T, Binks K, Tawn EJ, Burn J. 2004. Low erythrocyte folate status and polymorphic variation in folate-related genes are associated with risk of neural tube defect pregnancy. Mol Genet Metab, 81 :273-281 >DOI >Link

Wilding CS, Relton CL, Sutton MJ, Jonas PA, Lynch S-A, Tawn EJ, Burn J. 2004. Thymidylate synthase repeat polymorphisms and risk of neural tube defects in a population from the northern United Kingdom. Birth Defects Res A Clin Mol Teratol, 70 :483-485 >DOI >Link

Relton CL, Wilding CS, Jonas PA, Lynch SA, Tawn EJ, Burn J. 2003. Genetic susceptibility to neural tube defect pregnancy varies with offspring phenotype. Clin Genet, 64 :424-428 >Link

Wilding CS, Grahame J, Mill PJ. 2002. A GTT microsatellite repeat motif and differentiation between morphological forms of Littorina saxatilis: Speciation in progress? Marine Ecology Progress Series, 227 :195-204

Backeljau T, Frias Martins AM, Gosling E, Grahame J, Mill P, Brito C, Clarke A, Medeiros R, Small M, Wilding CS, Wilson I, Winnepenninckx B, Clarke R, de Wolf H. 2001. Periwinkles (Gastropoda, Littorinidae) as a model for studying patterns and dynamics of marine biodiversity Bulletin de L'Institut Royal des Sciences Naturelles de Belgique, Biologie, 71 :43-65

Mill PJ, Clarke AP, Smith DC, Grahame J, Wilding CS. 2001. Lagoonal littorinids: Shell shape and speciation Journal of Shellfish Research, 20 :469-475

Wilding CS, Butlin RK, Grahame J. 2001. Differential gene exchange between parapatric morphs of Littorina saxatilis detected using AFLP markers Journal of Evolutionary Biology, 14 :611-619 >DOI

Wilding CS, Grahame J, Mill PJ. 2001. Correlation of morphological diversity with molecular marker diversity in the rough periwinkle Littorina saxatilis (Olivi) Journal of Shellfish Research, 20 :501-508

Wilding CS, Grahame J, Mill PJ. 2000. Nuclear DNA restriction site polymorphisms and the phylogeny and population structure of an intertidal snail species complex (Littorina). Hereditas, 133 :9-18 >Link

Wilding CS, Grahame J, Mill PJ. 2000. Mitochondrial DNA CoI haplotype variation in sibling species of rough periwinkles. Heredity (Edinb), 85 ( Pt 1) :62-74 >Link

Wilding CS, Mill PJ, Grahame J. 1999. Partial sequence of the mitochondrial genome of Littorina saxatilis: relevance to gastropod phylogenetics. J Mol Evol, 48 :348-359 >Link

Simpson RJ, Wilding CS, Grahame J. 1999. Simple protocol for extracting nuclear DNA from single embryos of a marine snail. Biotechniques, 26 :1050-1052 >Link

Wilding CS, Beaumont AR, Latchford JW. 1999. Are Pecten maximus and Pecten jacobaeus different species? Journal of the Marine Biological Association of the United Kingdom, 79 :949-952 >DOI

Wilding CS, Grahame J, Mill PJ. 1998. Rough periwinkle polymorphism on the east coast of Yorkshire: Comparison of RAPD-DNA data with morphotype Hydrobiologia, 378 :71-78

Wilding CS, Latchford JW, Beaumont AR. 1998. An investigation of possible stock structure in Pecten maximus (L.) Using multivariate morphometrics, allozyme electrophoresis and mitochondrial DNA polymerase chain reaction-restriction fragment length polymorphism Journal of Shellfish Research, 17 :131-139

Wilding CS, Beaumont AR, Latchford JW. 1997. Mitochondrial DNA variation in the scallop Pecten maximus (L) assessed by a PCR-RFLP method HEREDITY, 79 :178-189 >DOI >Link

Wilding CS, Beaumont AR, Latchford JW. 1997. Mitochondrial DNA variation in the scallop Pecten maximus (L.) assessed by a PCR-RFLP method. Heredity (Edinb), 79 ( Pt 2) :178-189 >Link

Engagement & Impact

External PGR examinations performed:

Name of University: University of Liverpool, Award student examined for: PhD, Thesis title: A genomic approach to understanding metabolic insecticide resistance to diamides in diamondback moth (Plutella xylostella)

External PGR Supervision - completed students:

Name of University: University of Liverpool, Award gained by the student: PhD, Thesis title: Evolutionary genetics of insecticide resistance in Culex quinquefasciatus

Other invited event:

Title of event: Wales Gene Park Schools’ Genetics Roadshow, Location: Coleg Menai, Ffriddoedd Road, Bangor, Gwynedd, Description: Invited talk "The genomics of insecticide resistance in mosquitoes"

Title of event: Genetics for Teachers: CPD event, Location: The Racecourse, Wrexham, Description: Invited talk: "Unravelling species barriers – classroom applications of third generation DNA sequencing"

Research Grants Awarded:

Wellcome Trust, Wellcome Trust Biomedical Vacation Scholarship: "Cloning and sequencing of a carboxylesterase implicated in insecticide resistance in the malaria vector Anopheles arabiensis", Mr Daniel Kaptijn (LJMU Undergraduate), Grant value (£): 2000, Duration of research project: 8 weeks

Wellcome Trust, Wellcome Trust Biomedical Vacation Scholarship: "Development of novel molecular markers for the study of population structure in sub-Saharan African populations of Simulium blackflies, vectors of onchocerciasis", Ms. Calista Dufton-Kelly, LJMU undergraduate, Grant value (£): 2000, Duration of research project: 8 weeks

Teaching qualification:

Title of qualification gained: Fellow of the Higher Education Academy

Title of qualification gained: Staff and Educational Development Association: Professional Certificate in Supporting Learning