J Mallet CV 1

James Mallet Curriculum Vitae

31 October 2009

Personal Details

Email address: j.mallet@ucl.ac.uk Telephone: USA (617)-496-2780 Sabbatical 2009-2010:

Present appointment:

Radcliffe Institute of Advanced Studies Harvard University Byerly Hall 8 Garden St CAMBRIDGE MA 02138 Professor of Biological Diversity Department of Genetics Evolution and Environment University College London 4 Stephenson Way LONDON NW1 2HE

Education/Qualifications

1984 PhD Zoology University of Texas at Austin 1978 MSc Applied Entomology University of Newcastle-upon-Tyne 1976 BA Zoology University of Oxford Past Professional History

1996 - 2000 Reader in Evolutionary Biology, Department of Biology, University College London

1991 - 1996 Lecturer and Departmental Tutor, Galton Laboratory, Department of Genetics and Biometry, University College London

1988 - 1992 Assistant Professor, Insect Genetics, Department of Entomology, Mississippi State University

1985 - 1988 Postdoctoral Research Fellowship, Natural Environment Research Council, University College London

J Mallet CV 2

Other Appointments and Affiliations (since 1995)

2009 - 2010 Helen Putnam Fellow, Radcliffe Institute for Advanced Study, Harvard University

2008 - 2009 Fellow, Wissenschaftskolleg zu Berlin 2007 - 2009 Director, Centre for Ecology and Evolution 2007 Visiting Professor, Université de Perpignan (2 mo., June & October) 2007 - Guest Professor, Center for Computational and Evolutionary Biology

(CCEB), Beijing, China 2002 - Honorary Research Fellow, The Natural History Museum 2001 - 2004 Special Editor, "Evolution" Journal 2000 - Research Associate, Smithsonian Tropical Research Institute, Balboa,

Panama 1999 - 2006 Editorial Board, Proceedings and Philosophical Transactions of the

Royal Society of London 1999 - 2002 Research Associate, Department of Entomology, The Natural History

Museum, London (2002 - , Honorary Research Fellow) 1996 - 2000 Associate Editor, "Evolution" journal 1995 - 2005 Charles Darwin Trust, advisory role with, via Stephen Keynes, Chairman 1990 - Profesor Asociado, Instituto de Genética, Universidad de Los Andes,

Bogotá, Colombia 1987 - Investigador Asociado, Departamento de Entomología, Museo de

Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Perú

Prizes, Awards and other Honours

2009 Darwin-Wallace Medal, The Linnean Society of London 2009 - 2010 Helen Putnam Fellow, Radcliffe Institute for Advanced Study, Harvard

University 2008 - 2009 Fellow, Wissenschaftskolleg zu Berlin 2007 - Elected Fellow, the Linnean Society of London (FLS) 2002 - Elected Fellow, American Association for the Advancement of Science

(AAAS) 2002 - Elected Fellow, Royal Entomological Society (FRES) 2000 Vice President, Society for the Study of Evolution 1985 - 1988 Postdoctoral Research Fellowship, Natural Environment Research

Council of Great Britain 1973 - 1976 Frazer Scholarship, Balliol College, Oxford

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Grants (since 1995):

2009 - 2012 BBSRC Research Grant. Genomic analysis of complex speciation. £667,135 FEC UCL component. £850,000 total 2008 - 2011 Member of Leverhulme International Network: "Questioning the tree of life." PI John Dupré, University of Exeter (£71,619 among many international philosophers & historians of science) 2008 NERC Molecular Genetics Facilities Grant. SNP discovery in the

transcriptome of Heliconius. £3,000 2007 - 2012 University of London grant to the Centre for Ecology and Evolution.

London Evolutionary Research Network. £100,000 2006 - 2009 DEFRA Darwin Initiative. Tropical Andean butterfly diversity project.

£149,187 2004 - 2006 GBIF-ECAT grant. Global butterfly names. $US 50,000. 2004 - 2007 NERC Research Grant. Repeated genetic patterns across a faunal

"suture zone": £244,279 2004 - 2007 NERC Research Equipment. Sequencing and Genotyping to establish

The Centre for Comparative Genomics: £306,387 2003 - 2004 Natural History Museum Special Funds. Global butterfly nomenclature

database: £6,775. 2001 - 2003 EC-Marie Curie Postdoctoral Fellowship (Sebastien Gourbière).

€114,072 2002 - 2004 EC-Marie Curie Postdoctoral Fellowship (Jesús Mavárez). €114,072 2002 - 2004 EC-Marie Curie Postdoctoral Fellowship (Mathieu Joron). €114,072 2002 - 2004 EC-Marie Curie Postdoctoral Fellowship (Marie Zimmermann).

€114,072 2001 - 2004 Leverhulme Grant. Can mimicry cause speciation? £101,808 1999 - 2002 BBSRC Research Grant, Genomic mapping of reproductive isolation in

the larch budmoth: £208,216 1998 - 2001 NERC Research Grant. Selection and the nature of the species

boundary: £182,222 1998 - 2002 NERC - EDGE Research Grant. Genetic structure and spatial dynamics

of endemic butterfly races (jointly with Univ. Leeds and Univ. Birmingham): £325,240

1997 British Council Grant, travel to Colombia to teach on international course on phylogenetic methods in systematics and biogeography: £800

1996 Royal Society Conference Grant, for International Congress of Entomology, Florence: £565

1995 - 1998 BBSRC Research Grant, Horizontal gene transfer between biotypes of the larch budmoth, Zeiraphera diniana: £192,176

1994 - 1995 AFRC Research Grant, Genetics of demographically unstable insects: £59,712

1993 - 1997 SERC Research Grant, Genetics of speciation in a tropical butterfly: £181,584

1992 - 1996 NERC Research Grant, Genetic effects of population fragmentation in endangered British butterflies: £141,327

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Invited lectures (since 1995)

2009 "150 Years of Evolution. Darwin's Impact on the Humanities." College of Arts and Letters, San Diego State University

"Darwin's origin of species - in the light of modern research," 51st Phylogenetisches Symposium, Staatliches Naturhistorisches Museum, Braunschweig (turned down)

Radcliffe Institute for Advanced Study, Harvard University "Hybridization in Primates. Evidence, Extent, Evolutionary Impact and

Problems." Göttingen, Germany (turned down). "Perspectives on the Tree of Life." Philosophy, King's College University,

Halifax, Nova Scotia "Hybrid Speciation." European Society of Evolutionary Biology, Turin

(invited symposium organizer) "On the Origin of Species: Speciation Studies Now," Cambridge Darwin

Festival, Cambridge "Origin of Species–150 Years Later," Kristineberg Marine Research Station,

Sweden Staatliches Museum für Tierkunde, Dresden Royal Entomological Society (invited symposium organizer), "Insect

Evolution: Below the Species Level." Rothamsted, Harpenden, Herts Keynote Lecture, Deutsche Gesellschaft für allgemeine und angewandte

Entomologie, Göttingen, Germany Department d'ecologie et évolution, Université de Lausanne, Switzerland "Darwin Year 2009: Seminars on Evolution," Zoologisches Institut,

Universität Zürich, Switzerland University of Exeter at Cornwall, Plymouth 2008 Museum für Naturkunde, Humboldt-Universität, Berlin, Germany Institut für Populationsgenetik, Universität Wien, Austria Wissenschaftskolleg zu Berlin, Germany Border Natural History Society, Bellingham, Northumberland "Ecological Genomics of Speciation" Symposium, Ecological Society of

Japan, Fukuoka, Japan "Dynamic Interaction of Insects: Species, Colony and Individuals"

Symposium, Kobe University, Kobe, Japan ESRC Centre for Genomics in Society (EGENIS), University of Exeter International workshop: "Natural Selection and Speciation," Johannes

Gutenberg Universität, Mainz, Germany 2007 Department of Biological and Chemical Sciences, Queen Mary University of

London Centre for Population Biology, Imperial College (Silwood Park, Ascot) British Ecological Association. Education session. Sheffield University British Ecological Association. "Speciation and Ecology." Sheffield

University Hybridization and the Nature of Species, "The Legacy of Charles Darwin"

lecture series, Institute of Advanced Study, Durham University Symposium organizer and speaker. Evolution of Heliconius. International

Biology of Butterflies Conference, Rome

J Mallet CV 5

Invited symposium speaker: "Speciation and Natural Enemies," European Society for Systematic and Evolutionary Biology, Uppsala

Max Planck Institute for Chemical Ecology, Jena, Germany Speciation symposium, Systematics Association Meeting, Edinburgh

Plenary Speaker, 100th Meeting of the DZG (German Zoological Society), Cologne, Germany

Invited speaker: International Symposium on Phylogeography and Conservation, Luxembourg

2006 Conference on Biodiversity and Evolution, Mathematical Biology Institute, Ohio State University, Columbus, Ohio

Department of Zoology, University of Florida, Gainesville Department of Biological Sciences, University of Maryland, College Park "Darwin and 20th Century Science," Charles Darwin Trust, Seminar at

Trinity College, Cambridge "Integrating Evolution, Development and Genomics," Center for Integrative

Genomics, Integrative Biology and Molecular and Cell Biology, University of California, Berkeley

"A Century of Evolution: Ernst Mayr 1904-2005." Linnean Society, London "Parasite species and speciation – tackling a host of problems." Linnean

Society, London 2005 BBSRC Comparative Development (CODE) Initiative Workshop, Edinburgh

"Evolution and Development of Colour Pattern," Society for the Study of Evolution, Fairbanks, Alaska

Plenary speaker, “Bloodbath” Symposium, University of Stockholm, Sweden

Plenary talk, Department of Biology Graduate Symposium, Univ. Amsterdam, Holland

2004 Department of Biological Sciences, University of Hull The Royal Institution, London, schools talk: Sex in butterflies

"Evolution and Development in the Lepidoptera." International Conference, San Juan, Puerto Rico

Organizer Mimicry Symposium, International Congress of Entomology, Brisbane

University of Texas at Austin, Austin, Texas University of Florida, Gainesville, Florida (declined) "Architecture and Speciation", Architectural Association, London Plenary Talk, Finnish Graduate School in Evolutionary Ecology, University

of Jyväskylä, Finland 2003 Department of Genetics, University of Nottingham Department of Zoology, University of York

Ecological Genetics Symposium, International Congress of Genetics, Sydney, Australia

Evolutionary Genetics Symposium, Royal Entomological Society Annual Meeting, Reading (turned down due to other commitments, but coauthored chapter in symposium volume with Dr. Chris Jiggins)

Tropical Biology Association, Kibale Forest Reserve, Uganda (International graduate field course)

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London Evolution Research Network (London-wide postgraduate organization)

Division of Environmental and Evolutionary Biology, University of Glasgow Speciation Symposium, The Natural History Museum, London Department of Zoology, University of Cambridge Department of Entomology, The Natural History Museum, London

2002 Biology of Butterflies Conference. University of Leiden, Holland Department of Biology, University of Amsterdam (turned down) International Congress of Systematic and Evolutionary Biology. Butterfly

Biodiversity Symposium; from Genomics to Ecosystems. Patras, Greece (turned down)

Second International Heliconius Conference. Gamboa, Panama Imperial College London at Silwood Park, Ascot 2001 Institute of Zoology, London Entomology Department, Rothamsted Experimental Station, Herts Nils von Höfsten Memorial Lecture, Evolutionary Biology Centre, University

of Uppsala, Sweden Department of Psychology, University of Newcastle-upon-Tyne Institute of Cell, Animal and Population Biology, University of Edinburgh 2000 Aposematism: past, present and future. Symposium and Workshop.

University of Jyväskylä, Finland Survival of Species in Fragmented Landscapes. EU-funded Symposium.

Université de Montpellier II, France Department of Biology, University of Leiden, Netherlands Department of Biological Sciences, University of Sussex, Brighton Organizer, Evolution in Heliconius butterflies. Symposium of the

Association of Tropical Biology, Bloomington, Indiana CNRS, Université de Montpellier II, France Department of Biology, University of Stockholm, Sweden 1999 School of Animal and Microbial Sciences, University of Reading School of Biological Sciences, Royal Holloway and Bedford New College,

Egham Department of Biology, University of Maryland Institut de Zoologie, Université de Neuchâtel, Switzerland Royal Entomological Society Symposium on Insect Movement, London Department of Organismal and Evolutionary Biology, Harvard University Colour Group (UK arts/sciences interdisciplinary society), Royal Academy

of Arts, London 1998 Third International Butterfly Ecology and Evolution Symposium, Crested

Butte, Colorado (turned down) Department of Biology, University of Bristol (turned down) Department of Biology, University of Plymouth Institut des Sciences de l'Evolution, Université de Montpellier II, France

International Congress of Ecology, Florence, Italy; Symposium on Hybrid Zones (turned down)

Department of Zoology, University of Oxford Centre for Ecology and Evolution Symposium: Origins and Maintenance of

Biodiversity, Institute of Zoology, London

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Smithsonian Tropical Research Institute, Ciudad Panamá, Panamá 1997 Curso-Taller en Sistemática Filogenética, Biogeografía y sus Aplicaciones

en Biodiversidad y Conservación, Instituto de Ciencias Naturales, Villa de Leyva, Colombia

Instituto de Genética, Universidad de los Andes, Bogotá, Colombia Imperial College at Silwood Park, Ascot 1996 Setting Conservation Priorities, Centre for Conservation Genetics, Latrobe

University, Melbourne (paper presented in absentia) Department of Biological Sciences, University of Wales, Bangor "Endless Forms. Species and Speciation" meeting, Asilomar, California Hybrid Zones Symposium, International Congress of Entomology, Florence Genetic Conservation, The Mason Conference, British Association for the

Advancement of Science, Birmingham Department of Biology, Lund University, Sweden School of Biological Sciences, Queen Mary & Westfield College, London. 1995 Jacques Monod Conference on Evolution and Adaptation, CNRS, Aussois,

France Institut des Sciences de l'Evolution, Université de Montpellier, France Royal Society Meeting on Island Biology, London CIBA Foundation Meeting on Speciation, London

Academic supervision (since 1995): Postdoctoral researchers

2004 - Dr. Kanchon Dasmahapatra (funded by NERC, CEE, BBSRC): Repeated genetic patterns across a faunal "suture zone", followed by: Genomic analysis of complex speciation

2004 - 2008 Prof. Gerardo Lamas-Müller (funded by GBIF and DEFRA-Darwin Initiative): Butterfly nomenclature: Global Butterfly Names and Tropical Andean Butterfly Diversity projects

2005 - 2006 Dr. Vladimir Lukhtanov (funded by the Royal Society): European butterfly names

2002 - 2004 Dr. Mathieu Joron (EU Marie Curie Postdoctoral Fellow): The genetic architecture of colour pattern evolution in Heliconius butterflies. Currently a EMBL-funded postdoctoral fellow, University of Edinburgh. Will soon take up a permanent CNRS position at the Musée d'Histoire Naturelle in Paris

2002 - 2004 Dr. Marie Zimmermann (EU Marie Curie Postdoctoral Fellow): Phylogeny, speciation, mimicry, phylogeography of Melinaea. Currently lecturer at Université Saint Jerome, Marseille

2002 - 2004 Dr. Jesús Mavárez (EU Marie Curie Postdoctoral Fellow): Genetic analysis of speciation in Heliconius butterflies. Currently Assistant Professor, IVIC, Venezuela

2001 - 2004 Dr. Keith Willmott (funded by a Leverhulme grant): Can mimicry cause speciation? Currently Assistant Professor and Curator, Florida Museum of Natural History, Gainesville, Florida

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2001 - 2003 Dr. Sebastien Gourbière (EU Marie Curie Postdoctoral Fellow): Speciation theory: bridging the gap between ecological and genetic approaches. Currently a lecturer at the Université de Perpignan

1999 - 2002 Dr. Ian Wynne (funded by NERC - EDGE): Genetics of endemic butterfly races. Went onto an EU-funded postdoctoral position at University of Copenhagen

1998 - 2001 Dr. Chris Jiggins (funded by NERC): Selection and the nature of the species boundary. Currently Royal Society University Research Fellow, University of Cambridge

1994 - 1997 Dr. W. Owen McMillan (funded by BBSRC): Genetics of speciation in a tropical butterfly. Currently Full Professor, Univ. of Puerto Rico

1993 - 2001 Dr. Igor Emelianov (funded by Society for the Protection of Science and Learning, AFRC, UCL bridging funds, and BBSRC): Genomic mapping of reproductive isolation. Currently a tenured government research entomologist at IACR-Rothamsted

1993 - 1996 Dr. Martin Brookes (funded by NERC): Genetic effects of population fragmentation in endangered British butterflies. Currently a freelance journalist

Other salaried research assistants

2005 - 2008 Blanca Huertas (UCL/NHM, funded by DEFRA-Darwin Initiative as

database coordinator) 2005 - Mari-Wyn Burley (UCL, funded by NERC/UCL as technician) 2000 - Fraser S. Simpson (UCL, funded by UCL/NERC/BBSRC as technician) 1993 - 1997 Peter King (UCL, funded by UCL/BBSRC as technician) 1992 - 1998 Yvonne Graneau (UCL, funded by UCL/NERC/BBSRC as technician)

Postgraduate students i. Supervised

2008 - PhD supervisor for Neil Rosser (funded by NERC): Climate change and

biodiversity distribution in the tropics. UCL 2006 - PhD supervisor for Blanca Huertas (coordinator of DEFRA-Darwin

Initiative grant "Tropical Andean Butterfly Diversity Project"): Biodiversity of butterflies in Colombia. UCL/NHM

2006 - 2007 PhD supervisor for Jae-Woo Chung (overseas student funded by Kwanjeong Educational Foundation, Seoul, and Overseas Research Student [ORS] UK govt. award): Mate choice and speciation in Heliconius butterflies. UCL. (withdrew)

2005 - PhD supervisor for Lisa Leadbeater (funded by NERC): Hybridization and evolution in the genus Oleria (Ithomiinae). UCL

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2001 - 2009 PhD supervisor for Sarah Darwin (funded by BBSRC, interrupted several times due to repeated child-bearing): Systematics and evolution of tomatoes on the Galápagos Islands (Solanum, Solanaceae). UCL, awarded subject to minor corrections

2001 - 2004 PhD supervisor for Margarita Beltrán (overseas student funded by Overseas Research Student (ORS) UK govt. award, in collaboration with Smithsonian Tropical Research Institute (STRI), Panama): A genomic approach to speciation in the genus Heliconius. UCL. Awarded. Currently a secondary school science teacher

2001 - 2005 PhD supervisor for Alaine Whinnett (funded by NERC-CASE, in collaboration with the Natural History Museum): Molecular evolution in the Ithomiinae. UCL. Awarded. Currently a science teacher

1999 - 2003 PhD supervisor for Vanessa Bull (funded by NERC): Population genetics of Lepidoptera. University of London. Awarded. Currently working as a forensic scientist in the Forensic Science Centre, London

1997 - 2001 PhD supervisor for Russell Naisbit (funded by BBSRC): Speciation and hybridisation between Heliconius cydno and H. melpomene. University of London. Awarded. Currently a postdoctoral researcher at Université de Neuchâtel

1996 - 2000 PhD supervisor for Michele Drès (funded by NERC): Gene flow between host races of a forest insect. University of London. Awarded. Currently working as an NHS Administrator

1994 - 1997 PhD supervisor for Chris Jiggins (funded by BBSRC): The ecology and genetics of speciation in Heliconius butterflies (Lepidoptera: Nymphalidae). University of London. Awarded. Currently a Royal Society URF with a large group in Cambridge University

1993 - 1997 PhD supervisor for Owen Rose (funded by demonstratorship at UCL): "The evolution of microsatellites". University of London. Awarded. Currently in Business Education

In addition I have co-supervised 9 external UCL students under the London-based PRI scheme, and have acted as second supervisor for a number of internal UCL students

ii. Postgraduate examining

2008 PhD external examiner for Inke van der Sluijs: Divergent mating preferences and nuptial coloration in sibling species of cichlid fish. University of Leiden, Netherlands

PhD external examiner for Nicola Chamberlain: Molecular basis of Heliconius wing patterns. University of Exeter at Cornwall, Falmouth, Cornwall

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2007 PhD internal examiner for Paris Veltsos: Sex chromosome fusion and rDNA diversity in the grasshopper Podisma pedestris hybrid zone. Queen Mary University of London

2006 PhD external examiner for Thibaut Malausa: Isolement reproducteur, différenciation génétique et adaptation aux plantes hôtes chez deux espèces de lépidoptères du genre Ostrinia: un cas de spéciation écologique? Université Paul Sabatier, Toulouse

PhD external examiner for Siu Fai Lee: Genome mapping in Lepidoptera. University of Melbourne, Australia

2005 PhD internal examiner for Antonio Hernandez Lopez: Evolutionary dynamics of host plant use and diversification in two insect-plant interactions. Imperial College, University of London

2004 PhD internal examiner for Shen-Horn Yen: Phylogenetic reconstruction of the Chalcosiinae (Lepidoptera: Zygaenidae sensu lato). Imperial College, University of London

2003 PhD internal examiner for Nick Isaac: Continuous characters in macroevolution: testing with sister-clade comparisons. Imperial College, University of London

2000 PhD internal examiner for James J. Groombridge: Conservation genetics of the Mauritius kestrel, pink pigeon and echo parakeet. Queen Mary College, University of London

PhD external examiner for Cock van Oosterhout: Inbreeding depression and genetic load in laboratory metapopulations of the butterfly Bicyclus anynana. University of Leiden, Netherlands

PhD external examiner for Gabriella Gamberale-Stille: Behaviour and aposematism. University of Stockholm, Sweden

1999 - 2003 External Examiner, MSc Course: Advanced Methods in Taxonomy and Biodiversity. Imperial College/Natural History Museum, London

1999 PhD internal examiner for Jan H. Kolaczinski: Aspects of pyrethroid resistance in relation to impregnated bednets. London School of Hygiene and Tropical Medicine

1998 PhD external examiner for Angel Viloria: Systematics and evolution of high elevation Pronophilini (Lepidoptera: Satyrinae). Imperial College, University of London

PhD external examiner for Jonathan Bridle: Evolutionary genetics of Chorthippus grasshoppers in Europe. University of Leeds

1997 PhD external examiner for Carlos Pinheiro: Unpalatability, mimicry and escaping ability in neotropical butterflies: experiments with wild predators. University of Oxford

PhD external examiner for Durrell Kapan: Selection for mimetic polymorphism in the butterfly Heliconius cydno. University of British Columbia

pre 1997 I was internal examiner for 4 earlier University of London PhD theses

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James Mallet: Outline of research Current research interests a) Comparative genomics, especially to investigate adaptive evolution, genetics of species traits such as evo-devo of colour pattern development, as a means to study gene flow between species, and the nature of species and speciation. b) Genetics of species and speciation. Genealogical and coalescent approaches. c) Hybridization and hybrid speciation, and their importance in nature. d) Highly replicated biogeographic analyses in high-species diversity faunas as a comparative approach to study species diversification in the Amazon basin. e) Evolution and systematics of butterflies (especially Heliconiinae and Ithomiinae). f) Natural history, biodiversity and taxonomy informatics. g) History and philosophy of science (when necessary).

Research findings a) Selection on mimicry genes is strong, and important in the lives of Heliconius. b) Mimicry shifts cause or are associated with speciation in Heliconius. Mimicry shifts cause reproductive isolation as a by-product via two mechanisms: (a) strong selection against non-mimetic hybrids between Müllerian mimicry forms, and (b) pleiotropic effects of colour pattern on visually based mate choice. c) Mimicry differences between species (i.e. between H. cydno and H. melpomene, and between H. erato and H. himera) are inherited as single genes of large effect, which are the same as those known from geographic races within species. d) In work with Henri Descimon, I have shown that at least 16% of European butterfly species hybridize in nature with one or more other species. In another study, on wild-caught hybrids between species of Heliconius, I show that a large fraction (35%) of species hybridize. In about half of the cases, clear evidence for backcrossing exists, suggesting that genes may flow between species in nature. Genomic evidence proves that genes are regularly, albeit rarely introgressing among Heliconius species in nature, and may contribute to adaptive evolution of mimicry and other ecologically important traits. There is preliminary evidence for quite substantial amounts of 'hybrid speciation' and transfer of adaptive mimicry genes among members of the melpomene group of Heliconius. e) Gene sequencing studies have shown that H. cydno and H. melpomene have fixed differences at mitochondrial and some nuclear gene loci, but that the same species share identical haplotypes at other nuclear loci. Coalescent analyses of

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population sequence data implicate introgression between species and/or hybrid speciation as the cause. The genomes in these species therefore show preliminary evidence for 'islands of speciation' which determine species phenotypes, separated by regions that introgress more freely. f) The switch-supergene that controls polymorphic mimicry by H. numata of its ithomiine co-mimics is controlled by one of the same regions for mimicry in H. melpomene. We have sequenced the region on BAC clones in both species. Chromosomal arrangements are largely identical between species within Heliconius, and several similar-acting mimicry loci in H .erato/himera map to the same co-linear genomic locations in H. melpomene/cydno. These mimicry loci are therefore clearly conserved throughout Heliconius, but also have extremely flexible gene action, since the colour patterns of H. numata are radically different to those in H. melpomene or H. erato (in collaboration with M. Joron and Heliconius consortium). g) The rates of evolution of new geographic races and species in replicate Ithomiinae and Heliconius taxa across a narrow ‘suture zone’ in Peru are highly, and significantly variable. While some groups of species show very slow rates of speciation and mimicry evolution (e.g. Oleria), others (e.g. Melinaea) show extremely rapid rates. These results, obtained via novel multilocus coalescent analyses, cast doubt on the idea that mimicry evolution and speciation was driven simply by allopatry, for example after formation of Pleistocene forest refugia. Instead, biological, lineage-dependent differences among clades in response to the environment are most likely the cause of these variable rates of diversification. Simple ‘environmental forcing’ of species diversification does not seem a globally tenable explanation, given such widely differing rates of evolution in a single region. h) With a team of taxonomists, especially Gerardo Lamas and Keith Willmott, we have compiled a databased checklist of scientific names, including all subspecies and valid synonyms, as well as many invalid names of all butterflies on the planet, and type specimen photos and data for the neotropics for release on the internet. Linked to this project, we have museum and genetic/genomic specimen databases for the neotropical area, closing gaps between museum taxonomy and genomics informatics. i) Darwin's statements about species and speciation are widely misunderstood. Many historians of science have been aware of this, but even they have interpreted Darwin in the light of a perceived 'correct' reproductive isolation or phylogenetic view of species, developed in the 20th Century. In the 150th anniversary of Darwin's 'Origin', I have contributed strongly to a renewed assessment of Darwin's view of species. Erroneous heredity aside, Darwin's views are increasingly useful for interpreting today's molecular and genomic data on speciation.

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Achievements and influence

I have over 160 publications (excluding meeting abstracts, since first publication as an undergraduate in 1976), and I am known world-wide in evolution and genetics circles, leading to many invitations to speak at international meetings, especially on the topic of speciation. My associates, former associates and I have built Heliconius into a key organism for understanding evolutionary genetics, evolution of development, and speciation. Initial work was on hybrid zones between mimetic geographic races of H. melpomene and H. erato, leading to an understanding of mimetic selection pressure and gene flow. Recent work has concentrated on speciation and hybridization studies among full species, especially between H. himera and H. erato, and then H. cydno and H. melpomene, as well as polymorphic mimicry in H. numata. In the Heliconius melpomene group on which we are focusing new genomics approaches, we have preliminary evidence for hybridization and adaptive introgression among around 15 sympatric species. This work is leading to a revolutionary understanding of the nature of species, as well as providing mapping tools for identification and association mapping of 'speciation genes' and the evo-devo of colour pattern development. PhD students and postdocs from my group have now founded major laboratories of their own in Puerto Rico and North Carolina State (W. Owen McMillan), Cambridge (Chris Jiggins), and Edinburgh & Paris (Mathieu Joron). They form a large fraction of today’s international Heliconius Consortium (www.heliconius.org) involved in Heliconius ecology, behaviour, evo-devo and genomics research. Meanwhile, we have recently spearheaded genetic work to study evolution and mimicry of a different and somewhat lesser known group, the ithomiine butterflies, important Müllerian mimicry partners of some Heliconius. The great advantage of this group is replicability in a comparative approach – there are over 300 species, as compared with only ~40 in Heliconius. Ithomiine research has been carried out not only in my own laboratory, but in collaboration with Edinburgh (Marianne Elias, Mathieu Joron), University of Florida (Keith Willmott, another former postdoc), Cambridge (Chris Jiggins), and Lima (Gerardo Lamas).

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James Mallet: Teaching interests and other university service Overview and philosophy

I attempt to take fresh approaches in many problems in population genetics, evolution and biodiversity, and I find this attitude can be an exciting way to teach as well. I enjoy teaching, especially for the occasional successes one achieves as judged by comments from former students. I also like teaching because it forces me to think through topics related to, but outside my immediate specialisms. I am quite messianic that my undergraduates should learn enough maths and stats to understand basic population genetic and evolutionary theories, and analyse simple genetic data sets. This can be initially unpopular among the more innumerate undergraduates, but even the most disadvantaged usually agree afterwards that it is worthwhile. In my evolutionary genetics course at UCL, the math blow becomes softened as we get to more complex problems, such as sexual selection, speciation, and macroevolution: the teaching becomes more 'organismic' due to extreme complexity of mathematical treatment that would otherwise be required.

Potential courses and past teaching experience I currently teach an undergraduate course in evolutionary genetics, as well as a field course in ecological genetics which we hold in Southern Spain during the Mediterranean spring, and a biodiversity unit within a general ecology course (see: http://www.ucl.ac.uk/~ucbhdjm/courses). These receive excellent ratings at UCL. I would be happy to teach in a relatively broad range of undergraduate courses ranging from genetics, population genetics, evolution, population biology, to biodiversity etc., depending on Departmental teaching needs. While graduate courses are virtually non-existent in UK universities, I teach a one-day graduate course on likelihood and statistical inference with my colleague Ziheng Yang for the Graduate School at UCL. Future graduate courses could include areas of evolutionary genetics, comparative genomics, population genetics of biodiversity, etc. I also teach CoMPLEX graduate students ('enabling,' below). University service and 'enabling' At UCL I was until recently Director of the London-wide Centre for Ecology and Evolution (http://www.ucl.ac.uk/~ucbtcee/). I run the Centre for Comparative Genomics (and also was the PI on the NERC Equipment Grant that originally funded this service, see http://www.ucl.ac.uk/biology/centre-for-comparative-genomics/). I am a founding member of UCL's Centre for Mathematics and Physics in the Life Sciences and Experimental Biology (CoMPLEX, www.ucl.ac.uk/complex). CoMPLEX is UCL's interdisciplinary and inter-Faculty MRes/PhD graduate programme. Among many other minor jobs, I also redesigned and ran the entire main UCL Biology website (www.ucl.ac.uk/biology/) for many years.

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James Mallet: Publications (since 1995)

For electronic copies of many recent publications and papers in review etc., see: www.ucl.ac.uk/taxome/jim/jimpubs.html; for older publications, see: http://www.ucl.ac.uk/taxome/jim/dmp/jimallpubs.html For ISI disambiguation, e.g. from Jacques Mallet of human genome fame, see also my ResearcherID.com site: http://www.researcherid.com/rid/B-5114-2008

i. Books Mallet, J. (~2010). Evolution of Biological Diversity. Consequences of a Porous Species Boundary. (Proposal and sample chapter accepted for University of Chicago Press; this approx. 300 pp. book will cover the current revolution in understanding diversification and speciation).

ii. Edited books Patterson, C. (1998). Evolution. 2nd Edition. ix+166 pp. Natural History Museum/Cornell University Press. (Updated and edited by J. Mallet & P.L. Forey; foreword by P.L. Forey and J. Mallet). Loxdale, H., Claridge, M., & Mallet, J. (eds.) (2010). Darwin-Wallace Special Issue. Ecological Entomology (in press). iii. Book chapters Descimon, H. & Mallet, J. (2009). Bad species. In: Ecology of Butterflies in Europe (eds. Settele, J., Konvicka, M., Shreeve, T., Dennis, R. & Van Dyck, H.). Cambridge University Press. (in press). Mallet, J. (2009) Introduction. In Chan, C.-L. and G. W. Beccaloni (eds) Alfred Russel Wallace (1865). "On the Phenomena of Variation and Geographical Distribution as illustrated by the Papilionidae of the Malayan Region." Reprint edition. Natural History Publications (Borneo), Kota-Kinabalu, Malaysia (in press). Mallet, J. (2009) Rapid speciation, hybridization and adaptive radiation in the Heliconius melpomene group. In Butlin, R. Bridle, J. & Schluter, D. Speciation and ecology, Oxford University Press. pp. 177-194. http://www.ucl.ac.uk/taxome/jim/pap/Mallet_HeliconiusSpeciation.pdf. Mallet, J. (2008) Wallace and the agreed species concept of the early Darwinians. In Smith, C.R. and Beccaloni, G.W. (eds.): Natural Selection and Beyond: The Intellectual Legacy of Alfred Russell Wallace, Oxford University Press. pp. 102-113. http://www.ucl.ac.uk/taxome/jim/pap/Mallet on Wallace for Smith 2008.pdf

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Mallet, J. (2007). Species, concepts of. In Levin, S. et al. (eds.) Encyclopedia of Biodiversity. 2nd Edition. Elsevier. pp. 427-440. Mallet, J. (2007). Subspecies, semispecies, superspecies. In Levin, S. et al. (eds.) Encyclopedia of Biodiversity. 2nd Edition. Academic Press. pp. 523-526. Mallet, J. (2006). Species concepts. In: Evolutionary Genetics: Concepts and Case Studies (Eds. Fox, C.W. & Wolf, J.B.). Oxford University Press, Oxford, pp. 367-373. Jiggins, C.D., Emelianov, I. & Mallet, J. (2005). Assortative mating and speciation as pleiotropic effects of ecological adaptation: examples in moths and butterflies. In: Fellowes, M.D.E., Holloway, G., Rolf, J. (eds.) Evolutionary Ecology of Insects. Royal Entomological Society, London, pp. 451-473. Taneyhill, D.E., Mallet, J., Wynne, I., Burke, S., Pullin, A.S., Wilson, R.J., Butlin, R.K., Hatcher, M.J., Shorrocks, B., & Thomas, C.D. (2003). Estimating rates of gene flow in endemic butterfly races: the effect of metapopulation dynamics. In: Genes in the Environment. (Eds: Hails, R.; Beringer, J. & Godfray, H.C.J.). Blackwell Science, Oxford, 3-25. Mallet, J. (2001). Species, concepts of. In Levin, S. et al. (eds.) Encyclopedia of Biodiversity. Volume 5. Academic Press. pp. 427-440. Mallet, J. (2001). Subspecies, semispecies, superspecies. In Levin, S. et al. (eds.) Encyclopedia of Biodiversity. Volume 5. Academic Press. pp. 523-526. Mallet, J. (2001). Gene flow. In: Woiwod, I.P., Reynolds, D.R. & Thomas, C.D. (eds.) Insect Movement: Mechanisms and Consequences. CAB International, Wallingford, UK, pp. 337-360. Mallet, J., Jiggins, C.D., & McMillan, W.O. (1998). Mimicry and warning colour at the boundary between races and species. In Howard, D.J. & Berlocher, S.H. (eds.) Endless Forms: Species and Speciation. Oxford University Press. pp.390-403. Mallet, J.L.B., & Turner, J.R.G. (1998). Biotic drift or the shifting balance: did forest islands drive the diversity of warningly coloured butterflies? In Grant, P.R. (ed.) Evolution on Islands. Oxford University Press. pp. 262-280. Mallet, J. (1998). Species concepts. In Calow, P. (ed.) Encyclopaedia of Ecology and Environmental Management. Blackwell Press. pp. 709-711. Mallet, J. (1998). Isolating mechanisms. In Calow, P. (ed.) Encyclopaedia of Ecology and Environmental Management. Blackwell Press. pp. 379-380. Brown, T.M., Bryson, P.K., Arnette, F., Roof, M., Mallet, J.L.B., Graves, J.B., & Nemec, S.J. (1996). Surveillance of resistant acetylcholinesterase in Heliothis virescens. In Brown, T.M. (ed.) Molecular Genetics and Evolution of Pesticide Resistance. ACS Symposium 645: 149-157.

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Mallet, J. (1996). The genetics of biological diversity: from varieties to species. In Gaston, K. (ed.): Biodiversity: Biology of Numbers and Difference. Blackwell Press. pp. 13-47. Brookes, M., Emelianov, I., McMillan, O., Rose, O., & Mallet, J. (1996). A quick guide to genetic and molecular markers used in biodiversity studies. In Gaston, K. (ed.): Biodiversity: Biology of Numbers and Difference. Blackwell Press. pp. 48-53. iv. Articles in peer-reviewed journals de Silva, D.L., Silva, A., Gallusser, S. & Mallet, J. (2010) Selection for enemy-free space: oviposition away from the host increases survival in a tropical Ithomiinae butterfly. (manuscript in preparation) Dasmahapatra, K.K., Lamas, G., Simpson, F. & Mallet, J. (2010) Genetics and evolution of Amazonian butterflies: coalescent analyses show that speciation rates depend more on biological and lineage effects than on vicariance. (manuscript in preparation) Hill, R.I., Elias, M., Dasmahapatra, K., Willmott, K., Jiggins, C., Koong, V., & Mallet, J. (2009). Cryptic, ecologically differentiated species and polyphyly in the Mechanitis mazaeus complex (Ithomiinae). (manuscript in preparation) Mallet, J. (2009). Shift happens! Shifting balance and the evolution of warning colour and mimicry diversity. Ecological Entomology (in press). Baxter, S.W., Nadeau, N., Maroja, L., Wilkinson, P., Counterman, B.A., Dawson, A., Beltrán, M., Perez-Espona, S., Chamberlain, N., Ferguson, L., Clark, R., Davidson, C., Glithero, R., Mallet, J., McMillan, W.O., Kronforst, M., Joron, M., ffrench-Constant, R. & Jiggins, C.D. Genomic hotspots for adaptation: the population genetics of Müllerian mimicry in the Heliconius melpomene clade. PLoS Genetics (accepted). Mallet, J., Wynne, I.R., Thomas, C.D. (2009). Introgression, species limits and conservation implications of climate change: brown argus butterflies in Britain (Polyommatus subgenus Aricia). Insect Conservation and Biodiversity (submitted).

de Silva, D.L., Day, J.J., Elias, M., Willmott, K., Whinnett, A. & Mallet, J. (2009). Molecular phylogenetics of the neotropical butterfly tribe Oleriini (Nymphalidae: Ithomiinae) inferred from multiple gene sequences suggest introgressive hybridization between genera. Molecular Phylogenetics and Evolution (submitted). http://www.ucl.ac.uk/taxome/jim/pap/de Silva et al Oleria 09.pdf

Mallet, J., Meyer, A., Nosil, P. and Feder, J. L. (2009). Space, sympatry and speciation. Journal of Evolutionary Biology 22: 2332–2341.

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Mallet, J. (2009). Why was Darwin's view of species rejected, and how does it fare today? Biology and Philosophy (in press).

Mallet, J. (2009). Group selection and the biological species concept. Philosophical Transactions of the Royal Society (in press).

Mallet, J. (2009). Alfred Russel Wallace and the Darwinian species concept: his paper on the swallowtail butterflies (Papilionidae) of 1865. Gayana (in press).

Sachs, J.D., Baillie, J.E.M., Sutherland, W.J., Armsworth, P.R., Ash,N., Beddington, J., Blackburn, T.M., Collen, B., Gardiner, B., Gaston, K.J., Godfray, H.C.J., Green, R.E., Harvey, P.H., House, B., Knapp, S., Kümpel, N.F., Macdonald, D.W., Mace, G.M., Mallet, J., Matthews, A., May, R.M., Petchey, O., Purvis, A., Roe, D., Safi , K., Turner, K., Walpole, M., Watson, R. & Jones, K.E. (2009). Policy forum: Biodiversity conservation and the Millennium Development Goals. Science 325: 1502-1503.

Gourbière, S. & Mallet, J. (2009). Are species real? The shape of the species boundary with exponential failure, reinforcement, and the 'missing snowball.' Evolution (in press).

Dasmahapatra, K.K., Elias, M., Hill, R.I., Hoffman, J.I., & Mallet, J. (2009). Mitochondrial DNA barcoding detects some species that are real, and some that are not. Molecular Ecology Resources (available online). Mallet, J. (2008). Hybridization, ecological races, and the nature of species: empirical evidence for the ease of speciation. Philosophical Transactions of the Royal Society B – Biological Sciences 363: 2971-2986. Mallet, J. (2008). Mayr's view of Darwin: was Darwin wrong about speciation? Biological Journal of the Linnean Society 95: 3-16. Mallet, J., Beltrán, M., Neukirchen, W., & Linares, M. (2007). Natural hybridization in heliconiine butterflies: the species boundary as a continuum. BMC Evolutionary Biology 7:11. Beltrán, M., Jiggins, C.D., Brower, A.V.Z., Bermingham, E., & Mallet, J. (2007). Do pollen feeding and pupal-mating have a single origin in Heliconius? Inferences from multilocus sequence data. Biological Journal of the Linnean Society 97: 243-252. Mallet, J. (2007). Hybrid speciation. Nature 446: 279-283. Anstead, J.A., Mallet, J., and Denholm, I. (2007). Temporal and spatial incidence of alleles conferring knockdown resistance to pyrethroids in the peach-potato aphid Myzus persicae Sulzer, and their association with other insecticide resistance mechanisms Bulletin of Entomological Research 97: 243-252.. Elias, M., Hill, R.I., Willmott, K., Dasmahapatra, K.K., Brower, A.V.Z., Mallet, J., & Jiggins, C.D. (2007). Limited success rate of DNA barcoding in identifying

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butterflies from a diverse tropical community. Proceedings of the Royal Society B 274: 2881-2889. Dasmahapatra, K. K., Silva, A., Chung, J.-W., & Mallet, J. (2007). Genetic analysis of a wild-caught hybrid between non-sister Heliconius butterfly species. Biology Letters 3: 660-663. Mallet, J. (2006). What has Drosophila genetics revealed about speciation? Trends in Ecology and Evolution 21(7): 186-193. Joron, M., Papa, R., Beltrán, M., Chamberlain, N., Mavárez, J., Baxter, S., Abanto, M., Bermingham, E., Humphray, S.J., Rogers, J., Beasley, H., Barlow, K., ffrench-Constant, R.H., Mallet, J., McMillan, W.O., & Jiggins, C.D. (2006). A conserved genetic locus controls pattern diversity in Heliconius. PLoS Biology 4(10): e303. Bull, V., Beltrán, M., Jiggins, C.D., McMillan, W.O., Bermingham, E. & Mallet, J. (2006). Polyphyly and gene flow between non-sibling Heliconius species. BMC Biology 4:11. Dasmahapatra, K.K. & Mallet, J. (2006). DNA barcodes: recent successes and future prospects. Heredity 97: 254-255. Lamas, G. & Mallet, J.L.B. (2005). Case 3320. Papilio sapho Drury, 1782 (currently Heliconius sapho; Insecta, Lepidoptera): proposed conservation of the specific name. Bulletin of Zoological Nomenclature 62(1): 21-24. Mallet, J., Isaac, N., & Mace, G. (2005). Taxonomic inflation: reply (correspondence). Trends in Ecology and Evolution 20: 8-9. Mallet, J. (2005). Hybridization as an invasion of the genome. Trends in Ecology and Evolution 20: 229-237. Isaac, N.J.B., Mace, G.M., & Mallet, J. (2005) Response to Agapow and Sluys: the reality of taxonomic change (correspondence). Trends in Ecology and Evolution 20: 280-281. Gourbière, S. & Mallet, J. (2005). Has adaptive dynamics contributed to the understanding of adaptive and sympatric speciation? Journal of Evolutionary Biology 18: 1201-1204. Whinnett, A., Brower, A.V.Z., Lee, M.-M., Willmott, K.R., & Mallet, J. (2005). Phylogenetic utility of Tektin, a novel region for inferring systematic relationships amongst Lepidoptera. Annals of the Entomological Society of America 98: 873-886. Whinnett, A., Willmott, K.R., Brower, A.V.Z., Simpson, F., Lamas, G. & Mallet, J. (2005). Mitochondrial DNA provides an insight into the mechanisms driving diversification in the ithomiine butterfly Hyposcada anchiala (Lepidoptera: Nymphalidae, Ithomiinae). European Journal of Entomology 102: 633-639.

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Whinnett, A., Zimmermann, M., Willmott, K.R., Herrera, N., Mallarino, R., Simpson, F., Joron, M., Lamas, G. and Mallet, J. (2005). Strikingly variable divergence times inferred across an Amazonian butterfly 'suture zone'. Proceedings of the Royal Society B 272: 2525-2533.

Emelianov, I., Marec, F., & Mallet, J. (2004). Genomic evidence for divergence with gene flow in host races of the larch budmoth. Proceedings of the Royal Society of London B: 271: 97-105. Willmott, K.R. & Mallet, J. (2004). Correlations between adult mimicry and larval hostplants in ithomiine butterflies. Proceedings of the Royal Society of London B 271 (Biology Letters S5): S266-S269. Mallet, J. (2004). Poulton, Wallace and Jordan: how discoveries in Papilio butterflies initiated a new species concept 100 years ago. Systematics and Biodiversity 1: 441-452. Isaac, N.J.B., Mallet, J. & Mace, G.M. (2004). Taxonomic inflation: its influence on macroecology and conservation. Trends in Ecology and Evolution 19: 464-469. Mallet, J. (2004). Species problem solved 100 years ago (correspondence). Nature 430: 503. Emelianov, I., Simpson, F., Narang, P. & Mallet, J. (2003). Host choice promotes reproductive isolation between host races of the larch budmoth Zeiraphera diniana. Journal of Evolutionary Biology 16: 208-218. Mallet, J. & Willmott, K. (2003). Taxonomy: renaissance or Tower of Babel? Trends in Ecology and Evolution 18: 57-59. Knapp, S., & Mallet, J. (2003). Refuting refugia (commentary). Science 300: 71-72.

Naisbit, R.E., Jiggins, C.D. & Mallet, J. (2003). Mimicry: developmental genes that contribute to speciation [inheritance of mimicry differences between Heliconius cydno and Heliconius melpomene]. Evolution and Development 5: 269-280. Dasmahapatra, K.K., Blum, M., Aiello, A., Hackwell, S., Davies, N., Bermingham, E.P., Mallet, J. (2002). Inferences from a rapidly moving hybrid zone in sibling butterfly species (Anartia). Evolution 56: 741-753. Beltrán, M., Jiggins, C. D., Bull, V., Linares, M., McMillan, W.O., Mallet, J. & Bermingham, E. (2002). Phylogenetic discordance at the species boundary: gene genealogies in Heliconius butterflies. Molecular Biology and Evolution 19: 2176-2190.

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Drès, M. & Mallet, J. (2002). Host races in plant-feeding insects and their importance in sympatric speciation. Philosophical Transactions of the Royal Society of London B 357: 471-492. Naisbit, R.E., Jiggins, C.D., Linares, M., Salazar, C., & Mallet, J. (2002). Hybrid sterility, Haldane's rule, and speciation in Heliconius cydno and H. melpomene. Genetics 161: 1517-1526. Emelianov, I., Drès, M., Baltensweiler, W., Mallet, J. (2001). Host-induced assortative mating in host races of the larch budmoth. Evolution 55: 2002-2010. Jiggins, C.D., Naisbit, R.E., Coe, R.L., & Mallet, J. (2001). Reproductive isolation caused by colour pattern mimicry. Nature 411: 302-305. Mallet, J. (2001). Causes and consequences of a lack of coevolution in Müllerian mimicry. Evolutionary Ecology 13(7/8): 777-806. Joron, M., Wynne, I.R., Lamas, G. & Mallet, J. (2001). Variable selection and the coexistence of multiple mimetic forms of the butterfly Heliconius numata. Evolutionary Ecology 13(7/8): 721-754. Jiggins, C.D., Linares, M., Mallet, J., Naisbit, R.E., Salazar, C., Yang, Z. (2001). Sex-linked hybrid sterility in a butterfly. Evolution 55: 1631-1638. Mallet, J. (2001). Mimicry: an interface between psychology and evolution. (Commentary). Proceedings of the National Academy of Sciences USA 98: 8928-8930. Mallet, J. (2001). The speciation revolution (commentary). Journal of Evolutionary Biology 14: 887-888. Naisbit, R.E., Jiggins, C.D., and Mallet, J. (2001). Disruptive sexual selection against hybrids contributes to speciation between Heliconius cydno and H. melpomene. Proceedings of the Royal Society of London B 268: 1849-1854. Jiggins, C.D. & Mallet, J. (2000). Bimodal hybrid zones and speciation. Trends in Ecology and Evolution 15: 250-255. Jiggins, C.D., Mallet, J. (2000). Bimodal hybrid zones and the scale of a snail. (Correspondence: reply to M. Schilthuizen). Trends in Ecology and Evolution 15: 469. Bilton, D.T., Goode, D. & Mallet, J. (1999). Genetic differentiation and natural hybridisation between two morphological forms of the common woodlouse, Oniscus asellus Linnaeus, 1758. Heredity 82: 462-469.

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Mallet, J. & Joron, M. (1999). Evolution of diversity in warning colour and mimicry: polymorphisms, shifting balance, and speciation. Annual Review of Ecology and Systematics 30: 201-233. Joron, M. & Mallet, J. (1999). Diversity in mimicry (correspondence). Trends in Ecology and Evolution 14: 150. Davison, A., McMillan, W.O., Griffin, A.S., Jiggins, C.D., & Mallet, J.L.B. (1999). Behavioural and physiological adaptation between two parapatric Heliconius species (Lepidoptera: Nymphalidae). Biotropica 31: 661-668. Knapp, S. & Mallet, J. (1998). A new species of Passiflora (Passifloraceae) with notes on the natural history of its herbivore, Heliconius (Lepidoptera: Nymphalidae: Heliconiiti). Novon 8: 162-166. Mallet, J., McMillan, W.O., & Jiggins, C.D. (1998). Estimating the mating behavior of a pair of hybridizing Heliconius species in the wild. Evolution 52: 503-510. Joron, M. & Mallet, J. (1998). Diversity in mimicry: paradox or paradigm? Trends in Ecology and Evolution 13: 461-466. Jiggins, C.D., McMillan, W.O. & Mallet, J.L.B. (1997). Host plant adaptation has not played a rôle in the recent speciation of Heliconius himera and Heliconius erato (Lepidoptera: Nymphalidae). Ecological Entomology 22: 361-365. Brookes, M.I., Graneau, Y.A., King, P., Mallet, J.L.B., Rose, O.C., & Thomas, C.D. (1997). The genetic consequences of artificial introductions and natural colonizations in the British butterfly Plebejus argus. Conservation Biology 11: 648-661. Lewis, O.T., Thomas, C.D., Hill, J.K., Brookes, M.I., Crane, T.P.R., Graneau, Y.A., Mallet, J.L.B., & Rose, O.C. (1997). Three ways of assessing metapopulation structure in the butterfly, Plebejus argus. Ecological Entomology 22: 283-293. Davies, N., Aiello, A., Mallet, J., Pomiankowski, A. & Silberglied, R E. (1997). Speciation in two neotropical butterflies: extending Haldane's rule. Proceedings of the Royal Society of London B. 264: 845-851. McMillan, O., Jiggins, C., & Mallet, J. (1997). What initiates speciation in passion-vine butterflies? Proceedings of the National Academy of Sciences, USA 94: 8628-8633. Jiggins, C.D., McMillan, O., King, P. & Mallet, J. (1997). The maintenance of species differences across a Heliconius hybrid zone. Heredity 79: 495-505. Turner, J.R.G., & Mallet, J.L.B. (1996). Did forest islands drive the diversity of warningly coloured butterflies? Biotic drift and the shifting balance. Philosophical Transactions of the Royal Society (B) 351: 835-845.

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Mallet, J. (1996). What are good species? (Correspondence: reply to Shaw). Trends in Ecology and Evolution 10: 490-491. Mallet, J., Jiggins, C.D., & McMillan, W.O. (1996). Evolution: mimicry meets the mitochondrion. Current Biology 6: 937-940. Jiggins, C., Mallet, J., McMillan, O., & Neukirchen, W. (1996). What can hybrid zones tell us about speciation? The case of Heliconius himera and H. erato (Lepidoptera: Nymphalidae). Biological Journal of the Linnean Society 59: 221-242. Mallet, J., & Gilbert, L.E. (1995). Why are there so many mimicry rings? Correlations between habitat, behaviour and mimicry in Heliconius butterflies. Biological Journal of the Linnean Society 55: 159-180. Mallet, J. (1995). A species definition for the modern synthesis. Trends in Ecology and Evolution 10: 294-299. Mallet, J. (1995). A functional species definition. (Correspondence: reply to Gittenburger & Dover). Trends in Ecology and Evolution 10: 490-491. Emelianov, I., Mallet, J. & Baltensweiler, W. (1995). Genetic differentiation in the larch budmoth Zeiraphera diniana (Lepidoptera: Tortricidae): polymorphism, host races or sibling species? Heredity 75: 416-424.

v. Other publications Loxdale, H., Claridge, M., & Mallet, J. (2010). Preface. Darwin-Wallace Special Issue. Ecological Entomology (in press). Mallet. J. (2009). Ask the ecologist: Are genetic barcodes the magic tools we need to achieve a precise estimation of biodiversity in tropical ecosystems? What are the advantages and problems associated with this methodological proposal? Informative Bulletin of the Centro Internacional de Ecología Tropical (CIET), Caracas Vol. 1 (2): 3. Mallet, J. (2009). Darwin and Heliconius. heliconius.org. http://heliconius.zoo.cam.ac.uk/heliconius/2009/darwin-and-heliconius/ Mallet, J., Neukirchen, W. & Linares, M. (1997-2006). Hybrids between species of Heliconius and Eueides butterflies: a database. http://www.ucl.ac.uk/taxome/hyb/helichyb.html Mallet, J. (2005). Scaling problems in species diversity assessment. In: E-conference on Landscape scale biodiversity assessment: the problem of scaling. Centre for Ecology and Hydrology/European Platform for Biodiversity. http://www.edinburgh.ceh.ac.uk/biota/Archive_scaling/6721.htm

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Mallet, J. (2004). The peppered moth: a black and white story after all. Genetics Society News 50: 34-38. Also at anti-creationist site: http://www.talkreason.org/articles/mallet.cfm Mallet, J. (2004). Antenna: freely available online, and Poulton on species 1904. Antenna 28: 220-221. Mallet, J. & Thomas, C.D. (2003). 4.10 Brown argus butterfly. In Crane, P. & Bateson, P.: Measuring Biodiversity for Conservation. Policy Document 11/03. The Royal Society, London. pp. 36-38. Mallet, J. (2002). Memorandum by Professor James Mallet. In House of Lords: What on Earth? The Threat to the Science Underpinning Conservation. Stationery Office Ltd, London. Vol. 2, pp. 34-40. Mallet, J. (2000). Comments by James Mallet on A.R. Wallace (1864) "On the phenomena of variation and geographical distribution as illustrated by the Malayan Papilionidae". http://www.wku.edu/~smithch/S096.htm#"Mallet" Mallet, J. (2000). Comments by James Mallet on A.R. Wallace (1866) "Natural Selection". http://www.wku.edu/~smithch/S123.htm#"Mallet" vi. Book, film and other media reviews Mallet, J. (2008). The Grants of the Galapagos. Review of "How and Why Species Multiply: The Radiation of Darwin's Finches" by Peter R. Grant & B. Rosemary Grant. Trends in Ecology and Evolution 23: 535-536. Mallet, J. (2005). Speciation in the 21st Century. (Review of "Speciation" by Jerry A. Coyne & H. Allen Orr). Heredity 95: 105-109. Mallet, J. (2005). Biodiversity - the airbrushed version. (Review of: "Biodiversity: An Introduction", 2nd Edition, by K.J. Gaston and J.I. Spicer). Systematics and Biodiversity 3: 219. Knapp, S. & Mallet, J. (2005). A night out with the nerds. (Review of: “Theatre of Science,” a performance by Simon Singh and Richard Wiseman at the Soho Theatre, London) PLoS Biology 3(9): e325, pp. 1532-1533. Mallet, J. (2004). A butterfly book for mini-bucks. (Review of "Butterflies: Ecology and Evolution Taking Flight," ed. by C.L. Boggs, W.B. Watt & P.R. Ehrlich). Trends in Ecology and Evolution 19: 119-120. Mallet, J. (2004). Fritz Müller: A Naturalist in Brazil. By David West. Quarterly Review of Biology 79: 196-197.

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Mallet, J. (2004). From a ‘feel for the organism’ to a model system. (Review of "On the Wings of Checkerspots. A Model System for Population Biology" ed. by P.R. Ehrlich & I. Hanski). Trends in Ecology and Evolution 19: 625-626. Mallet, J. (2002). Move over Darwin. (Review of "In Darwin’s Shadow: The Life and Science of Alfred Russel Wallace" by Michael Shermer). Nature 419: 561-562. Mallet, J. (2001). Holy Landscapes! (Review of “Epistasis and the Evolutionary Process”, edited by Jason B. Wolf, Edmund D. Brodie III, and Michael J. Wade). Science 291: 602. Mallet, J. (2001). Taxonomy and the blues. (Review of “Nabokov’s Blues,” by Kurt Johnson and Steve Coates). Nature 409: 664-665. Mallet, J. (2001). Pioneers, dandies and frauds: 300 years of British butterfly collectors. (Review of “The Aurelian Legacy. British Butterflies and their Collectors”, by Michael A. Salmon). Endeavour 25: 135. Mallet, J. (2000). The new Lepidoptera systematics. (Review of “Handbuch der Zoologie. Band/Volume IV. Arthropoda: Insecta. Teilband/Part 35. Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics, and Biogeography.” Edited by Niels P. Kristensen. Berlin, Walter de Gruyter). Zoological Journal of the Linnean Society 129:267-269.

Mallet, J. (2000). Species and their names. (Review of “Species: New Interdisciplinary Essays,” edited by R.A. Wilson). Trends in Ecology and Evolution 15:344-345. Mallet, J. (2000). The gift of Lepidoptera. (Review of “Nabokov’s Butterflies: Unpublished and Uncollected Writings”, edited by Brian Boyd and Robert Michael Pyle). Nature 406: 827-828. Mallet, J. (1999). Review of “Molecular Genetic Analysis of Populations. A Practical Approach, 2nd Edition”, edited by A.R. Hoelzel. Annals of Human Genetics 63: 273-274. Mallet, J. (1998). Review of "Adaptation", edited by Michael R. Rose and George V. Lauder. Heredity 80: 393-394. Mallet, J. (1998). Demes, hosts and speciation. (Review of "Genetic Structure and Local Adaptation in Natural Insect Populations", edited by S. Mopper & S.Y. Strauss.) Ecological Entomology 23: 495-496. Mallet, J. (1998). Tropical ecology in miniature. (Review of "The Butterflies of Costa Rica and their Natural History. Volume II: Riodinidae", by P.J. DeVries). Trends in Ecology and Evolution 13:377.

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Mallet, J. (1997). Review of "Ecological and Evolutionary Aspects of Insecticide Resistance", by John A. McKenzie. Genetical Research 68: 183-187. Mallet, J. (1997). What are species? (Review of "Species, the Units of Biodiversity", edited by M.F. Claridge, H.A. Dawah, and M.R. Wilson.) Trends in Ecology and Evolution 12: 453-454. Mallet, J. (1996). Molecular evolution: selection wins the debate. (Review of "The Causes of Molecular Evolution", by J.H. Gillespie). Annals of Human Genetics. 60:85-86. Mallet, J. (1995). Review of "The Structure and Confirmation of Evolutionary Theory", by Elizabeth A. Lloyd. Heredity 75: 112. Mallet, J. (1995). Review of "Evolution", by Mark Ridley. Heredity 75: 223-224.