AGROPOLIS INTERNATIONAL agriculture • food • environment

Agropolis International Agropolis International is an international agricultural campus. brings institutions It represents a significant potential for scientific and technological of research and higher education capability: over 2,200 scientists in more than 110 research units in in Montpellier and Montpellier and Languedoc-Roussillon region including Languedoc-Roussillon 300 scientists in 60 countries. into partnerships with local communities, The scientific community focuses on the major issues related to : companies and enterprises in the • Agronomy in Mediterranean region, and into and tropical agricultural production sectors, close co-operation • Biotechnology and food technology, with international institutions. • Biodiversity, natural resources and ecosystems, • Water, environment and sustainable development, Agropolis International • Rural development and societies, is a world centre open to the stakeholders • Genomics in plant and integrative biology, involved in the • Food quality and safety. economic and social development in the following fields: Agropolis International is a centre for the capitalisation and agriculture, food, valorisation of knowledge, for training personnel and for technology environment transfer; it is a hub for visitors and international exchanges; it promotes actions based on multilateral expertise and contributes to the scientific and technological knowledge needed for preparing development policies. approach. Inagriculture, thechallenge • setupbiocontrol strategiesusingall biodiversity, andecology in agroecosystems andinnatural strategies, incorporatingthestudy of is tostudyandpreserve biodiversity • understandtherole ofbiodiversity research inMontpellier constraints influencingdevelopment preserved, conserved, restored, and complementary competenciessuch complementary are directly involved in research on population ecology andcomplexity,population ecology biological control aswell asseveral • accountfor physical andchemical that needstobeurgently analysed, French scientificcampusofitskind improved, allthrough asustainable Scientific activityby naturalistshas Within the Agropolis International Improvement ofbiologicalcontrol and one of the most important in and oneofthemostimportant in theequilibriumofecosystems; become amajorissueworldwide multidiscplinary skillsinorder to: multidiscplinary dynamics oflivingorganisms, and of organismsinecosystems; and Europe, dealingwithvarious and with regard totheenvironment. community, aboutfiftyscientists research concernsbiodiversity and theLanguedoc- • analyze therole oflandscape structures intheevolutionary One ofthemaingoals ofthis Roussilon region in • useappropriate agricultural in plantprotection This constitutesthepremier biological control, available naturalresources, Competencies in in thefollowing chapters. hundred more indirectly. practices for preventing from insectstoviruses; is realized today using managed landscapes. their interactions. as thosecovered pest outbreaks ;

biodiversity, andecology „ „ „ „ „ „ „ „ diversity anditsinteractions Understanding andevaluating in agroecosystems Characterising biodiversity Foreword List ofabbreviations Glossary at Agropolis International andtrainingEducation by research teams Topics covered scientific andtechnicalculture Dissemination of Controlling populations Characterization ofgeneticdiversity Evaluating hostspecificity Legislationandregulatory aspects... ofinvasive species Assessment Taxonomic tools Agroecology: anothervision Sustainable plantprotection Classicalbiologicalcontrol and interactions biologicalcontrolserving of sustainableagriculture for sustainableagriculture in plantprotection Biological control, Page 56 Page 44 Page 40 Page 26 Page 58 Page 55 Page 54 Page 52 Page 32 Page 22 Page 20 Page 12 Page 34 Page 24 summaryPage 8 Page 6 Page 4 4 Biological control species hadtakenadvantage biological control originated control ofsuchinvasive pest ecological, biotechnological, environments withouttheir agents), where theinvading of approaches builtaround and biodiversity. Biological of theirabsencetobecome or antagonists(biocontrol species covers amultitude an ecologicalequilibrium and geneticmanagement by introducing closelyco- evolved natural enemies as atechniquetorestore can threaten ecosystems introduced intoforeign native natural enemies Populations ofspecies options. Historically, invasive pests. inundative biologicalcontrol. that are oftenequallyconsidered as allelochemicals, natural toxins) ingredients (e.g. 2ºplantcompounds, to “natural” mixtures ofactive biopesticideisalsoapplied The term viruses, ornematodes.microsporidia, bacteria (e.g.bacteria including fungi(mycoherbicdes), and are basedonlivingagents phytopathogen orweed targets commercialised againstinsect, than 150biopesticideshavebeen and glasshouse. For example, more indigenous pestsinboththefield control in thelast30years totarget allowed inundativebiological or cosmopolitanantagonistshas outbreaks. Use ofeitherindigenous suppressionshort-term ofpest biopesticides withtheaimof and registered non-persistent or commercially developed using mass-reared antagonists control butisarepetitive approach parallels classical biological Inundative biologicalcontrol typically anon-commercial solution. inputsandassuchis with nofurther provides management permanent the developmentalstage, potentially species andwhilecostlyslow in to well-established exotic invasive biological control islargely restricted provide control. Classical longterm and suppressing pestpopulations to natural enemyestablishingitself effectiveness isdependentonthe into thepest’s invaded range and these antagonistsare thenreleased release. In theabsenceofsuchrisks, species intheproposed area of native orcommercially important toanyprospective non-targetrisks range, whichthenare screened for antagonists ofthepestfrom itsnative control relies onco-evolved,specific biological control. Classicalbiological “classical” and “inundative” B commonly splitinto invasive pestsismost iological control ofalien Bacillus thuringiensis Bacillus ), Foreword chemicals intheenvironment. are aimedatdecreasing theuseof where publicandgovernment goals area inapestmanagementworld biological control akeygrowth crops) can beincluded. This makes pest management(includingGM or mutualistaimedatimproving manipulation ofthecrop, antagonist Now almostanybreeding orgenetic definition ofbiologicalcontrol. broadening anddiversifyingthe Recent moleculartechnologiesare based research. within thedefinitionofbiocontrol- of crop-pest-antagonist interactions aspects oftheecologyorevolution developments havebrought most such beneficialspecies. These as foodsources andrefuges for include increasing biodiversity in thecrop ecosystem.Examples enemy populationsalready present or encourage orprotect natural or microclimate ormutualisms) pests (e.g. manipulatingthesoil of crops to counterattacksfrom activities thatimprove thecapacity pest management. This includes systems approaches are usedin has developedinrecent years as biologicalcontrolConservation persistence ontheaffectedcrop. simply pre-empt pestinfection or applying beneficalspeciesthatmay biological control caninclude peaks. Furthermore, inoculative suppression ofpestpopulation of peststoprovide short-term released intooutbreak populations natural enemiesare inoculatively indigenous pestswhere known native provides amore “classical” optionfor Inoculative biologicalcontrol Andy Sheppard (C.I.L.B.A) Podagrion pachymerum (, Torymidae), female layingeggs on anootheca (Walker) © G. Delvare

maculosa) (Centaurea knapweed plant aphids ona Colony of © R. Sforza

5 Biological control © R. Sforza Characterising biodiversity in agroecosystems

S Parcelled landscape The simple classification of plant and animal organisms at the integrating hedges and genus or species level, whether collected in a far away country borders with high plant or in France, is a challenge in itself. Identifying studied material is diversity in Languedoc- a key step for a successful biocontrol programme. This requires Roussillon. precise and important skills that are becoming increasingly rare in world research institutes and universities. The scientific community of Agropolis International can provide these skills for a number of taxonomic groups that play a role in applied and fundamental research. Biocontrol programmes benefit from this network to identify key pests and their associated natural enemies in a local, regional, national, E.U., and world regulatory framework. Biological control Biological

6 p. 20 quarantine issues agreements and organisms,harmful regulatory aspects: Legislation and p. 12 invasive species Considering p. 8 biological control serving Taxonomic tools © D. Coutinot © G. Delvare © R. Sforza community. followed by eachscentistbelongingto Agropolis International materials from onepointtoanotherisframedby officialrules quarantine facilities(for foreign material). This exchangeof reared outeitheringreenhouses (for French material)or end, naturalenemiesare takenand backtothelaboratory plants orinsectscollectinginnationalparks). Inthe for French territory, (e.g. ifnecessary inthecaseofprotected program. This hastobedoneinforeign countriesaswell as to collectinnaturalhabitatsare requested for eachresearch the enemiesofthosenaturalenemies. Officialauthorizations the trophic complexes ofthetarget, itsnaturalenemies, and type, plantassociations, inorder tobetterunderstand etc.) ecological traitsare reported(impactofnaturalenemies, soil fungi). Systematicsskillsare required here. Inaddition, many (e.g. , mites, andpathogens(e.g. etc.) bacteria, viruses, the targetspeciesinitsnative habitatwithitsassociatedfauna Field surveys constitutethesecondstep, collectingsamplesof target speciesisalsounderstudy: barcoding. new approach thatconsidersthegeneticsignature ofeach step fallsundertheumbrella ofclassicalsystematics, buta the target’s native andinvaded rangesare necessary. This first boatnists, entomologists, acarologists, andnematologistsfrom and insectcollectionconsultations, aswell ascontactswith its geographical distribution; literature survey, herbarium steps, thefirstofwhichisstudyingtargetspeciesand Research inbiocontrol follows aseriesofcoordinated only habitatdestruction. second-leading threat toimperiled speciesdiversity, trailing their ranges. Invasive speciesare now regarded asthe invasions must besoughtasinvasive speciesrapidly expand and naturalecosystems. Long-termsolutionstobiological plant trades), may have seriousconsequencesonmanaged introductions, oftenaccidental(viatourism, war, seedand is continuously risingonallseven continents. These by airandsea, thenumber ofnewly introduced species Due totheincrease ofworldwide commercial connections scientists. species are amajorresearch topicfor Agropolis International species negatively impactingagro-ecosystems. Invasive biopest conceptiscommonly usedfor indiginousorexotic several naturalenemiesinteractingwitheachother. The two-species complexoramultiple-species schemeincluding C a pestanditsnaturalenemies. Itmay involve a precisely definethecomplexrelationships between haracterising biodiversity inbiocontrol aimsto and Marie-Stéphane Tixier-Garcin (Montpellier SupAgro) René Sforza (EBCL)

7 Biological control 8 Biological control associated withthesepests will certain agriculturalcertain chemicals. favourable conditionsfortheir unintentional introduction of Large-scale changesinspecies Large-scale ecosystems. Systematics teams Mediterranean basin, finding systematics specialistiscalled For example, tropical disease quality control, toensure the potentially usefulbiocontrol increased pressure from pest The intensificationofglobal species islikelyduetorecent investigations oncomplexes based oncurrent andfuture vectors andcrop pestscould new speciesintonon-native have amajorrole toplayin involved lateratthelevel of control program toidentify biovigilance programs. The development there. Finally, of biologicalcontrol agents distributions are predicted identity ofreleased agents. commerce andtravel also become establishedinthe upon early inabiological invasive speciesandtheir restrictions ontheuseof increase inimportance. agents. He orsheisalso inevitably includesthe global climatetrends. Therefore, taxonomic serving biologicalcontrolserving developed includethoseforpests Biological modelsthathavebeen to revise taxonomic relationships. ofpopulations, or history evolutionary for classifications, tounderstandthe investigations, eithertogiveabasis They produce phylogenetic programs.implicated inconservation insects,important andorganisms see box) ofcrop pests, medically database («Barcoding » project, pests). They haveinitiatedamolecular quarantine organisms andmajor databases) forcrop pests(including They developinteractive tools(e.g. the developmentofdiagnosticaids. systematics andidentification research anddevelopmentthrough control projects. They support inbiological protection, particularly organisms thatare involvedinplant International campus istoidentify that are present ontheAgropolis The missionofthesystematicsteams hosts). coevolution ofparasites withtheir between organisms (forexample oftaxaandinteractions history understanding oftheevolutionary Phylogenetics alsoenablesabetter classification oflivingorganisms. Systematics isthescienceof divergent biologicalcharacteristics. complexes orpopulationswith ofspecies finediscrimination very morphometry, enable andkaryology studies. DNAsequencing, are establishedthrough phylogenetic ofsharedcriteria ascendanceand Present classificationsrely onthe classification oflivingorganisms. Systematics isthescienceof in Agropolis International Systematics Taxonomic tools documents publications andspecificteaching The results are madeavailable through bioindicators (neuropterans). (culicidmosquitoes)and importance (honeybee), insectsofmedical wasps (chalcidids),pollinators hyphomycetes, parasitic bacteria), agents (phytoseiidmites, lacewings, insects), predatory biocontrol (rodents, nematodes, mites, * Text by written Gérard Delvare (Cirad), [email protected] genetics. between taxonomy andpopulation andthelinks population interface entities andinvestigatethespecies- to optimize themanagementofthese areas andhelps ofgeneticpeculiarity CBGP projects. This research identifies species andstudiedinother and co-phylogeography ofendangered higher taxa,aswell asphylogeography phylogenies ofgroups ofspeciesand cytogenetic and/ormolecular research dealingwithmorphological, addition there ismore fundamental of ashared systematicsdatabase. In which benefitfrom thedevelopment and maintenanceofcollections, identification tools;anddevelopment of identificationkeys;development characterization ofspecies;production including revision, and description, taxonomic research, strictly concerns classified intwocategories. The first SupAgro, Cirad, IRD)canbe (« CBGP»UMR1062Inra, Montpellier Biologie desPopulations etdeGestion phylogenetics »ofthe on thetheme«systematicsand The studiesthatwere out carried scientists at«CBGP» and biocomputing Systematics scientists * . Centre de © R. Sforza contrasting biologicalcharacteristics, can reveal species complexes with characterization ofstudied species scientific results. Moreover, fine non research subjectsisa The accurate identificationof biology, ecology, andevolution. providing related information totheir of organisms orcommunitiesby • Contributing tothemanagement divergence oftaxa; mechanisms responsible forthe • Understanding theevolutionary of theecologicalphenomena; phylogenetic analysiswiththestudy the studiedtaxaby integrating relationshipsthe historical of • Proposing classificationsreflecting identification tools; anddetection diagnostic criteria characterize biodiversityandprovide barcoding database)identificationto and/or molecular(e.g PCR,SCAR, tools ofmorphological,cytogenetic, • Developing efficientandeffective newspecies; if needed,describing theirtaxonomic statusand, clarifying etc.)(control, ; conservation, • Differentiation ofmanagedspecies diseases. Its maingoalsinclude: agents, as well asvectorsofhuman of crop pests andotherbiocontrol biological modelsincludingpredators biocomputing teamworks with The CBGPsystematicsand conditionofthevalidity of sine qua needs. Organisms ofagronomic, out forresearch andprofessional and taxonomic studiesare carried mites. Diagnosis, identification, groupsor various ofinsectsor Research Area 1 (4) Integrated crop protection; ; inventories (3) Biodiversity, faunistics, morphological phylogeny; taxonomy ; (2)Molecular and (1) Diagnosis, identification, which are strongly linked: develops fourmainareas ofresearch et zoologie agricole The teamcalled« documantation » called«technical • Aplatform of theJoint Research Unit, biology » shared withtheotherteams called«Molecular• Aplatform electronic microscope », called«Scanning • Aplatform including about2millionspecimens named«collection» • Aplatform developed by «CBGP»: Several are technicalplatforms efforts. of conservation speciesthatareheritage thesubjects pests, vectorsofhumandiseases, or is represented by phytophagous of biologicaldiversity, whetherit modes andcostsofthemanagement and haveastrong impactonthe deals withone » ofthe«CBGP Écologie animale Collecting natural enemies, mainly insects, Psylloidea. Agromyzidae ;andHomoptera : Diptera the (theflies),inparticular :Phytoseiid mites ; expertise team hasacknowledged taxonomic economic importance, forwhichthis three groups ofmajor ofarthropods ones. Current research focuseson new classificationsandtoreview old these relationships istopropose investigation. The aimofstudying levelsof taxaatvarious various relationshipshistorical between and assessment ofevolutionary Research Area 2 «Molecularplatform biology». through collaboration withtheCBGP biology methodsandtechniques molecular it benefitsfrom modern of appliedentomology. Moreover, that isuniqueinFrance inthefield technicallibrary and mitesarich collectionsofinsects has important catalogs, anddatabases. This unit to developnewidentificationtools, species characterization andhelp They alsohighlightproblems of oftheterritory.biological supervision need forregular fieldscoutingand native, orintroduced pests, andthe problems duetonew,phytosanitary These studiesdraw attentiontorising areimportance mostlyconcerned. environmental, orheritage on leafy spurge intheSaône on leafyspurge Valley ••• deals withthe

9 Biological control 10 Biological control Eurytomidae): Eurytomidae): top oftheantenna; takenwithascanningelectronic Michel Martinez (Inra),Michel Martinez Jean-François Germain(LNPV), Philippe Reynaud (LNPV), Jean-Claude Streito (LNPV) Marie-Stéphane Tixier-Garcin (MontpellierSupAgro), Marie-Stéphane Tixier-Garcin (MontpellierSupAgro), (Inra), BrunoMichel(Cirad), FranckDorkfeld (Inra), 3. Brian Rector, William Meikle, DominiqueCoutinot, Naturelle, assignedby IRD), Armelle Cœur-d’Acier Systèmes Populations-Environnement » Johan Michaux(FNRS)GauthierDobigny (IRD), Gérard Delvare (Cirad), Mireille Fargette(IRD), Thematic «Systématique-Taxonomie, Laurent Granjon(MuseumNationald’Histoire Researchers involved:Researchers Team «Écologie animaleetZoologie Jean-Pierre Quéré(Inra), MariaNavajas (Inra), working onthistheme Diadromus collaris Researchers involved:Researchers Serge Morand(IRD),SylvieManguin(IRD), (Inra, MontpellierSupAgro, Cirad, IRD) Andy Sheppard (1 Directors andresearchers involved: Researchers involved:Researchers Team «Écologie intégrative des (Inra, MontpellierSupAgro, Cirad, IRD) (Hymenoptera, ): CBGP, Centre deBiologie et CBGP, Centre deBiologie et Phylogénie-Phylogéographie » 1. Serge Kreiter (MontpellierSupAgro), 2. de GestiondesPopulations de GestiondesPopulations Mic Julien (2 Team «Luttebiologique contre lesbio-invasions » contre les bio-invasions » Director: DenisBourguet, Team «Luttebiologique [email protected] [email protected] Aximopsissp. Director: Denis Bourguet, Laboratoire Européen Laboratoire européen Researchers - teachers involved: -teachers Researchers Eurytoma sp. Eurytoma [email protected] agricole - Acarologie » Director: Walker Jones, Main teams de LutteBiologique USDA-ARS, EBCL, Other teams [email protected] [email protected] Alain Migeon(Inra) Alain Migeon(Inra) frontal viewtaken withascanning parasitoid ofdiamondbackmoth; UMR1062 Franck Hérard UMR 1062 detail oftheultrastructure ofthe CSIRO, Walker Jones, RenéSforza, nd semester2006), Denis Bourguet(Inra), Maria Navajas (Inra), st electronic microscope, 70x. semester2006), (Hymenoptera, (Hymenoptera, microscope, 5000x. (Gravenhorst) adult female. Biological Control atAgropolis »(C.I.L.B.A.) The «InternationalComplexof 1 2 Available onwww.cilba.agropolis.fr Contacts: Andy Sheppard andRégisGoebel, [email protected] • leadingdiscussiongroups suchasthe Informal group ofconfinedstructures advising • organizationofinternshipsandscientificopendays onspecifictopics; inthesustainable development• participation ofenvironmental agroecosystems; • evaluation ofhow toreduce chemicalinputs; • development ofefficientstrategiestoreduce crop lossesandprotect natural Studies conductedwithintheofCI.L.B.A. framework include: taxonomy, molecularbiology, classicalbiologicalcontrol, ofinvasive andecology species. laboratories share multiple programs, skills, technologies, andknow-how regarding knowledge inIPMstrategiesfor crops, pastures, andnaturalecosystems. C.I.L.B.A. isaninternationalnetwork ofresearchC.I.L.B.A andeducationprofessionals improving programmes onaninternationalscale. and rearing ofsuitablenaturalenemiesandthedevelopment ofcrop protection population dynamics, genetics, andmicrobial epidemiology ecology, theselection This approach requires bothfundamentalandapplied research, on in particular weeds, pathogens, toprevent etc.) orlimittheirdamagenuisance value. strategies. Biologicalcontrol involves theuseofnaturalenemiespestspecies(insects, pest populationsincrop protection, health, andenvironment, usingbiologicalcontrol comprisesfiftyscientists whostudymechanismsandregulation processesC.I.L.B.A of biological control atthelocal, nationalandinternationallevel. USDA/ARS andSPV), aswell astodevelop commonperspectives in CNRS-CEFE, Cirad, IRD, Universities Montpellier2&3, CSIRO, EBCL organize relationships between itsmembers(Inra, MontpellierSupAgro, C.I.L.B.A, created in1989, of ispart Agropolis Internationalandaimsto different agriculturalinstitutesonsecurityandsafety. resources; Taxonomic tools

© G. Delvare © G. Delvare serving biologicalcontrolserving

© H.-P. Aberlenc 3 Barcoding: of datafrom reference specimensandtomaintain adatabaseof protected species, etc.). Itisalsoindispensabletoensure traceability legislation(regardingelement ofthepertinent quarantine organisms, using traditionalmorphologicalmethods. Infact, thisisarequired cimens thathave beenunequivocally identified by taxonomicexperts However, to establishreference itiscompulsory sequencesusingspe- highly efficienctfor theidentification oftaxawithdifficulttaxonomy. Molecular identification, whichisregularly usedinprokaryotes, is 7. validationofthe«Barcoding »approach ispossiblethrough trial 6. abilitytoassessthequalityandeven theprobability oferror inan 5. incomplicatedmorphological terminology notrainingnecessary 4. uniformity ofthemolecularmethodology, asopposedtomorpho- 3. species(i.e. detectionofcryptic thoseindistinguishableatthe 2. reliable identificationfrom allthestagesofdevelopment (eggs, 1. errors associatedwithphenotypicvariabilityare eliminated(relative taxonomic identificationinclude: reliable. Several argumentsfor usingmolecularcharactersfor seems possibletoimprove themethodinorder tomake itmore in thefieldsofagriculture, health, andconservation. Nevertheless, it gh currently controversial, willbeaccessibletonearly allscientists and itslow cost, suggestthatwithin10years thistypeoftool, althou- The firstresults, showing theaccuracyofidentification, itsspeed, of theintraspecificvariabilityaspecies. group ofsequencesthathasbeenpreviously definedasrepresentative the sequences, by computinggeneticdistance, orby assignmenttoa This globalapproach isbasedontheinvestigation of ofthehomology as «barcoding ». ItusestheInternettolinkwith«identifier». This identificationmethod, assisted by biomoleculartracersisknown the mitochondrialgene«COI»)hasrecently beenputforward. based onthediagnosticpower ofsmall genefragments(mainly using A new approach fororganisms, taxonomicidentificationofeukaryotic Characterising biodiversity and biologicalcontrol. phytophagous mites, theirpredators, products on effects ofphytosanitary an ecotoxicological studyofthe integrated crop protection, with to theimplementationof knowledge thatwillbeindispensable to betterscientificandtechnical Research Area 4 as well asabroad. research inFrance andinventories) these locations(through ecological order toassessthebiodiversity of in natural ordisturbedhabitatsin sampled identification ofarthropods Research Area 3 groups. application toalargenumber oftaxafrom distinctphylogenetic identification; dependingonthe taxon andthedevelopmentalthat canvary stage; for allthetaxonomicgroups; logical identification, whichrequires complex keys thatare different morphological level); larvae, etc.); reliability ofthesequences); should contribute shouldcontribute dealswiththe evolution or in agroecosystems and professional entomologists. French amateur, andinternational, relationships withalarge network of with itsmembers. It maintains see box herewith) andcollaborates International deLutte Biologique activities of«C.I.L.B.A. »( inthe This teamalsotakespart European firms. phytosanitary chambers ofagriculture, and agricultural teachingcolleges), many «CETA »institutes(technical Institutes (ACTA, ITV, CTIFL), of Plant Protection, the Technical and abroad includingalltheservices network ofcollaborators inFrance This teamworks withanextensive revolution indiagnosis? Contact: Jean-Yves Rasplus, [email protected] nematodes), aswell as vectors ofhumandisease. database ofcrop pestsandbiocontrol agents(e.g. insects, mites, the future CIRESare actively involved inthe buildingofsucha The systematicsandbio-computing researchers oftheCBGPand detection ofinvasive specieseasier andmore reliable. increasing samplesizesandinfraspecificvariability, thusmakingthe addition ofsuchnew information tocontinually updatethedatabase, specimen for additional studies. This userapproach alsoenables the occur inFrance, itwillthenbepossibletoutilizethesequenceofthis corresponds tosequencesofexoticspeciesthatare notknown to If thesequencethatissubmittednotfound inthedatabase, or if it trol willbeprovided withtheidentification. tion, bibliographic references, andeven recommendations for biocon- An outputform includinginformation onsynonyms, ecology, distribu- 3. The userwillreceive thenameofnearest taxonwhichisinthe 2. The submittedsequenceiscompared tosequencesthatare already 1. The qualityofthesequenceisassessedby checkingfor the obtained isanalyzed withrespect tothefollowing: receives inexchangeaspeciesidentification. The sequencethatis sequences obtainedinthisway intoadatabaseviatheInternetand ded from theInternet. This userthensubmitsoneorseveral DNA company, whowillf for any speciesofinterest by sendingaspecimentosequencing A userofthe«Barcode »systemcangetsequenceinformation crop pestsorofdiseasevectors andthuslimittheirimpacts. effectivebe aparticularly tooltorapidly detectnew introductions of are allthemore whenconsideringthat«Barcoding important »could the sequencedspecimeninataxonomiccollection. These capabilities small organismswithoutdestroying them, thusallowing retention of specimens. Current molecularmethodsenablethesequencingofvery integrated information accumulated from collectedandsequenced based ondivergence from thenearest sequence. database, withanestimationofthereliability ofthisidentification in thedatabase; highly preserved sites, andby countingambiguousnucleotides. presence orabsenceofastopcodon, theabsenceofmutation at Complexe , ollow astandard protocol thatcanbedownloa- operational in 2009. operational in2009. Protection). This Centre shouldbe and theNational ofPlant Laboratory Museum National d’Histoire Naturelle IRD, Cirad, Montpellier SupAgro, many actorsatthenationallevel(Inra, of together thetaxonomic expertise in Montferrier-sur-Lez (34)grouping « CBGP»ontheBailarguet campus created. It willbeintegrated within et d’expertise ensystématique Centre inter-organismes derecherche systematics anddiagnosis(CIRES: Soon acentre ofexcellence in insystematics expertise of research and Inter-institutional centre The «CIRES»project : ) willbe 11 Biological control 12 Biological control Assessment ofbioinvasions Assessment requires multidisciplinary with theinvasion process Tracing thegeographical invasive species, together reduction ofbiodiversity. is important inaglobal is important not, mayinduceserious Newly introduced alien context, especiallywith of habitats, including species, accidentalor scientists ofdifferent interaction between international trade. damage inalltypes the proliferation of movement ofthese approaches and backgrounds. with invasiveness? or ecologicalprocesses associated 3. What are thebiological traits and/ communities? structure anddynamicsofisolated consequences ofinvasions onthe 2. What are theecological successions andspeciesinteractions? of theinvasive speciesonnatural 1. impact What isthelongterm main questions: Studies are organized underthree community structure. interactions between speciesand in populationdynamicsand studies onregulatory mechanisms such disturbancesrequires specific communities andpopulationsto Predicting theresponse ofindigenous associations andcomplexes. foreign ecosystemscreates new introduction ofexotic speciesinto of indegenouspopulations. The worldwide, andthreaten theviability homogenization offlora andfauna ecosystems, tothe contribute the restoration ofdeteriorated Bioinvasions inhibit canseriously (processes andconsequences). Cirad) bioinvasions concerns UM2, UM3,Montpellier SupAgro, Évolutive, UMR5175-CNRS,UM1, (Centre d’Écologie Fonctionnelle et Systèmes Écologiques « department et activitéshumaines team « One ofthetopicsdevelopedby the consequences processes and Bioinvasions: Écologie despopulations Dynamique des Dynamique invasive species Assessment of »ofCEFE »inthe programs. the invasive weeds and insectpests and plantpathology, integrated within molecular biology, insectpathology, The mainunitsatEBCLinclude laboratory.and multidisciplinary students working inaninternational twenty employees, plusseveral studied. EBCLcomprises particularly biology andbioinvasion processes are as biologicalcontrol agents. Pest facilities andeventuallydeveloped inquarantineexperimentation are characterized through careful (insects, mitesandpathogens). These exploration tofindnatural enemies of classicalbiologicalcontrol is come intoplay. Onecornerstone botany, acarology, andecologyall disciplines suchasentomology, weeds andinsectpests. Scientific to reduce theimpactofinvasive technologies thatcanbeused develop classicalbiologicalcontrol of research atEBCLhasbeento From thebeginning,overall goal at Auch, France in1919. overseas established USDAlaboratory whose roots trace backtothefirst Eurasia are orAfrica studiedatEBCL, enemies. from Those originating introduced withouttheirnatural many non-nativepestshavebeen 1 ARSscientists. In theUnited States, built by ARSandstaffedby Category andtheonlyoneownedService) and Agriculture, Agricultural Research of of USDA-ARS(USDepartment is themainoverseas laboratory Biological Control (EBCL) Laboratory ofMontpellier,north theEuropean Located ontheBaillarguet Campus, bioinvasions intheU.S. in France combatting A USDAlaboratory ••• Will thetree ofheaven A highlyinvasive tree inEurasia the caseofcommonragweed Invasion, agriculture andpublichealth: threshold oftheallergicriskisonly 5grains/m July. Pollen production canreach three trilliongrainsperplantwhilethe Plan on Air Quality. Pollen isproduced ingreat quantitybeginningin mid- classified asabiologicalpollutantaccording totheRhône-AlpesRegional Pollen ofcommonragweed, whichishighly allergenicfor many people, is cultivation, establishmentofdensecover crops, andmowing. effective herbicideuse)andculturalpracticessuchashoeing, hand-pulling, ment practicesfor control ofambrosia includecrop rotation (toenable inadditionharvest toreducing yields. Current integratedweed manage- herbicides. Inpeaandsoybean, ambrosia cancausedifficultieswith close taxonomicrelationship tosunflower limitsthepossibilityofusing competitive,and very itcancausetotalcrop loss. However, ambrosia’s soybean, maize, andsorghum. Insunflower fields, where itisadventitious sector, commonragweed isaproblem incrops like sunflower, pea, se ofthehighallergeniccapacity ofitspollen. Intheagricultural for thefarmer, whoseyieldsare threatened, asfor societyatlargebecau- monocultures, commonragweed isbecomingatrue “plague” asmuch With populationsspreading quickly from afew scattered plantstodense thoroughfares, railroads, andinagriculturalriparianzones. degraded anddisturbedhabitats. InFrance, itisfound mainly alongmajor that colonizeabandonedarablelands, roadsides, wasteareas and other genic pollen. Commonragweed oftendominatesinpioneercommunities Europe andFrancepresents seriousproblems becauseofitshighly aller- valleys. Today itcanbefound allover France. Itsrapid spread through for many years inseveral French regions: intheRhôneandSaône starting After aninitialperiodofslow populationgrowth, ithasbeenspreading dermatitis, urticaria, andeczema. The financialimpact inRhone-Alpes logic report, CAREPS, 2000). The symptomsinclude rhinitis, conjunctivitis, from ragweed allergy, withapossiblerangeof3-20% (Source: epidemio- A studyinRhône-Alpesshowed that6-12%ofthepopulationsuffers introduced accidentally intoFrance, perhaps attheendof19 throughout theworld (Allard 1945inMcKone et Tonkyn, 1986). Itwas America. Itisnow «common»underallthetemperatelatitudes annual plantbelongingtothe Asteraceae family thatcomesfrom North Common ragweed, Ambrosia artemisiifolia (Ailanthus altissima) L., isamonoeciouswind-pollinated 3 . lead ustohell? th century. © R. Sforza population oftheperiodsgreatest risk(www.rnsa.fr nizations. A nationalaerobiological inspection network existsandinforms ofRoadEngineeringandMotorways,Department andvarious localorga- Ambrosias (AFEDA), theRegionalDirectorate oftheEnvironment, the many institutionswork, including: theFrench Association oftheStudy common ragweed posesabroad socio-economicproblem onwhich number, cost268200Euros alone. etc.) intheRhônedepartment Thus, campaign (hand-pullingcampaign, distribution ofbrochures, toll-free rers )whilein2003, (Source URCAM the “Ambrosia” public awareness 2004 was1,2millionEuros for thetreatment ofragweed suffe- allergy [email protected] Contacts: andCindy Arnaud Martin Adolphe, policy ofstudyandmanagement thisbiologicalinvasion. ture of “goodwill” would make itpossibletoestablishacoordinated creation struc- ofanindependentbodyand/orinterdepartmental by commonragweed makes itboth asocialandeconomicissue. The The combinationofagriculturaland publichealthproblems caused preventive orcurative controls according tothedegree ofinfestation. tive actionsofprivateindividualsaswell astechnicalpersonnel, and toactaccording tothegravitynecessary oftheproblem viapreven- andcoordinated managementplanisestablished.2) aconcerted Itis the biology, ecology, anddemographics ofthisinvasive speciesandif be carried outconcretely ismadetobetterknow only if1)aneffort In spiteofthisawareness, thecontrol ofthisspecieswillbeableto (Source: decree “Spécial Ambroisie” Isère). premises) toprevent thegrowth andspread ofambrosia - public domainsoftheState(e.g. routes, transportation working to theobligationimposedonlocalauthoritiesandmanagersof common ragweed must becarried outby theowner” inaddition ragweed grows” andthaton “cultivated land, thedestructionof common ragweed, cleanandmaintainallspaceswhere common stipulate thatcitizensmust: “prevent thegrowth ofseedlings Direction oftheMedicalandSocialBusinesses, respectively) Direction of Agriculture andtheForest orDepartmental Ambrosia” dependingontheinstitutioninvovled (Departmental Prefectoral decrees, suchas “Fallows Ambrosia” or “Special Inflorescence ofcommonragweed © A.Martin ). 13 Biological control 14 Biological control

© M. Volkovicth hazardous topublichealth: Feasibility ofabiologicalcontrol againstanexoticplant Team «Écologie intégrative desSystèmes Researchers involved:Researchers Brian Rector, William Meikle, DominiqueCoutinot, UM1, UM2, UM3, MontpellierSupAgro, Cirad) Thématique « Bioagresseurs émergents Thématique «Bioagresseurs Researchers involved:Researchers X Institut desSciencesdel’Évolution S [email protected] Director: Jean-Dominique Lebreton, Andy Sheppard (1 Directors andresearchers involved: Larva of Larva Researchers involved:Researchers (Inra, MontpellierSupAgro, Cirad, IRD) Adultof released on Populations-Environnement » Researcher involved:Researcher de GestiondesPopulations [email protected] Mic Julien (2 John Thompson, Anne Charpentier John Thompson,Anne fonctionnelle etévolutive CEFE, Centre d’écologie contre les bio-invasions » ...continued onpage 16 Director: DenisBourguet, Team «Luttebiologique [email protected] Laboratoire Européen Laboratoire européen Director: NicolePasteur, Centre deBiologie et [email protected] Director: Walker Jones, Main teams de LutteBiologique USDA-ARS, EBCL, UMR 1062CBGP, [email protected] et bio-invasions » [email protected] (UMR 5175, CNRS, Alain Migeon(Inra) Franck Hérard (UM2, CNRS) Zygogramma suturalis Zygogramma suturalis Arnaud Martin, Cindy Adolphe, CSIRO, Walker Jones, RenéSforza, nd semester2006), Ambrosia Maria Navajas (Inra), st Sandrine Maurice semester2006), in Russia of plantspecies beingtargeted for • invasive plantecology: distribution; along withstudiesoftheirbiology and are subjectedtohost-range studies in Australia. Selected potentialagents The maintarget organisms are pests insects, andotherinvasive species. • classicalbiologicalcontrol areas: capacity forresearch inthefollowing Australia. providesThe laboratory the for agriculture andtheenvironment in risks, andbiosecurity phytosanitary concentrated oninvasive species, The central themeofresearch is sustainable agricultural development. involvedin institutions andindustry research research withinternational exploration andcollaborative engagedinbiological is primarily organization. In Europe, CSIRO CSIRO, Australia’s leadingresearch is theEuropean focalpoint for The CSIRO European Laboratory in Australia to studythebio-invasions …and theCSIROunit Contact: DominiqueCoutinot, [email protected] would bedesirable. this plantisautopiangoal, aEuropean program onthefeasibility ofbiologicalcontrol does notcurrently seemtobeapopulargoal inFrance. Even ifthetotaleradicationof insects withintheframework ofaclassicalbiologicalcontrol program againstambrosia of agriculturalproduction andsuccessseemstobemoderate. The useofphytophagous vicinity wasnoted, butthedensity of Soviet Union. The eliminationoftheplantsatpointsrelease andtheimmediate Of these, to control Stobaera concinna of Georgia, Ukraine, Russia, inex. Yugoslavia, Chinaand Australia. Inaddition, Zygogramma suturalis Australia; in Russia; in Russia; T in Russia; in Australia andChina; countries: have beencarried outfor biologicalcontrol ofragweed invarious Releases ofphytophagous insectsoriginatinginNorth America and otherorganismsare being studiedfor theirpossibleuse. within theframework ofclassicalbiologicalcontrol againstambrosia its native rangeinNorth America. organismsare Certain already used insects) exert pressure oncommonragweed Many potentialbiologicalcontrol agents(e.g. fungi, rusts, phytophagous Zygogramma bicolorata Tarachidia candefacta Zygogramma suturalis Zygogramma disrupta Epiblema strenuana rigonorhinus tomentosus rigonorhinus Parthenium hysterophorus,Parthenium Ambrosia artemisiifolia (Stål)(Homoptera: Delphacidae)hasbeenreleased in Australia (Fabricius)(Coleoptera: inRepublic Chrysomelidae) Euaresta bella populations (Walker) (Lepidoptera: Tortricidae) (Hübner)(Lepidoptera: Noctuidae) Rogers(Coleoptera: inRussia; Chrysomelidae) isestablishedin16areas and4republics oftheformer of weeds, Pallister(Coleoptera: in Chrysomelidae) (Say) (Coleoptera: Anthribidae) another weed affecting publichealth. (Loew) (Diptera: Tephritidae) Z. suturalis Assessment of hemispheres. by andsouthern thenorthern of theopposing seasonsexperienced time ofresults by taking advantage horticulture, • counter-season research in introduced toAustralia; organisms willbeselectedto ensuresrisk thatonlyhighlyspecific control agents, theassessmentof plants inAustralia. For biological pests anddiseasestocrops andnative posedbyinvestigate therisk European assessment:• risk effective biologicalcontrol agents; range studiesandfortheselectionof basisforhost- anevolutionary forms of geographical phylogeniesalso region oforigin. The development target organismright anddefineits are routinely usedtoidentifythe populations: • geneticsofplantandpest the invasive mechanisms; environment (Australia) tounderstand environment (Europe) totheexotic The studiescompare thenative to reproduction andsurvivorship. evaluate the factorsthatcontribute biological control are studied to (Ambrosia artimisiifolia) remains low (0,2insects/m²) inzones shortening the delivery thedelivery shortening molecular techniques

invasive species researchers in © M. Volkovicth Characterising biodiversity in agroecosystems

Studies of program on invasive mites in Europe); An ongoing study of specimens from phytophagous mites and iii) Characterization of important around the world aims to precisely biological traits by applying a genomics trace the invasion history of T. evansi. Many species of approach to detect candidate genes. Besides providing information are important crop pests. The Studies on systematics, phylogeny and about the dissemination pathways management of these organisms phylogeography require molecular of the mite, the study of the genetic requires a better understanding of the tools which are developed in the diversity also strives to detect the area historical and ecological factors acting CBGP’s molecular biology platform. of origin of T. evansi, which will direct upon the structure and functioning the search for efficient, well-adapted of their populations (e.g. migrations, natural enemies to control this pest. outbreaks, gene flow). Among the Innovation Several examples of this approach members of the CBGP (UMR 1062 among mites: exist in entomology, but they are rare – Inra, Montpellier SupAgro, Cirad, in acarology. This study is conducted IRD), this approach is used by the genetic tracking at CBGP (team “Integrative ecology of team ‘Écologie intégrative des Systèmes population-environment systems’), Populations-Environnement ’ (group Study of an important mite crop in collaboration with the University ‘Evolutionary processes of pest pest, Tetranychus evansi, benefits of Sao Paulo, Brazil (Departamento populations’) to study crop pests as from the diverse expertise of CBGP de Zooloia, ESALQ). As an example, well as medically important insects. scientists. As a first step to any study, a previous study of another The studied topics are more specifically an unambiguous identification of tetranychid, the cassava green related to species complexes, as well as the species is needed. This task can mite, Mononychellus progressivus, invasions of new geographical regions be difficult for very small organisms which originated in South America and colonization of new habitats. like mites whose size is 250 to 500 and was introduced into Africa, Fundamental questions concerning microns. has contributed to integrated mite the evolutionary biology of the studied management strategies to control this species are also addressed. Research conducted by different pest in colonized regions. Agropolis International teams (CBGP, The team working on phytophagous ‘Integrative ecology of population- A second example is the coconut mites is specifically interested in: environment systems’ and ‘Animal mite, the eriophyid Aceria i) Phylogeography and population ecology and agricultural zoology guerreronis, an emerging pest of genetics (concerning invasive species, – Acarology’) employ complementary coconut palms. A genetic study based mainly crop pests such as Tetranychus approaches to better understand the on worldwide mite sampling and use urticae and T. evansi, see box on page geographical origin and the colonizing of molecular markers has validated 17); ii) Morphological and molecular paths followed by this invasive mite. the hypothesis of a New World origin systematics, phylogeny of species Goals include finding predatory of this mite (despite the probable complexes, and databases (e.g. online mites that can efficiently control the Asian origin of coconut). This is an database of the Tetranychidae family; pest and predicting and preventing important result affecting quarantine contribution to the CEE-DAISIE expansion of pest mite populations. measures, especially in Asia. •••

Invasion of water primrose in Southwestern France

© A. Dutartre Biological control Biological 15 16 Biological control Michel Martinez (Inra),Michel Martinez Jean-François Germain(LNPV), Philippe Reynaud (LNPV), Jean-Claude Streito (LNPV) Marie-Stéphane Tixier-Garcin (MontpellierSupAgro), tropical, Cirad/Université delaRéunion) team isnotspecialized inbiological ten PhD studentsandpostdocs. The scientists, 3techniciansandabout CNRS). This teamincludes10 de l’Evolution »(UMR5554UM2- belongs tothe«Institut desSciences etCoévolutionConservations » The team«Métapopulations, and genetics populations dynamics Using toolsof Marie-Stéphane Tixier-Garcin (MontpellierSupAgro), (Inra), BrunoMichel(Cirad), FranckDorkfeld (Inra), Team «Écologie intégrative desSystèmes Researchers involved:Researchers Naturelle, assignedby IRD), Armelle Cœur-d’Acier Johan Michaux(FNRS)GauthierDobigny (IRD), Team «Unitéd’évaluation génétique» végétaux etbioagresseurs en milieu Gérard Delvare (Cirad), Mireille Fargette(IRD), Thematic «Systématique-Taxonomie, Laurent Granjon(MuseumNationald’Histoire Team «Écologie animaleetZoologie Olivier Bonato(IRD), FrédéricPellegrin (IRD), Jean-Pierre Quéré(Inra), MariaNavajas (Inra), working onthistheme UMR C53PVBMT(Peuplements Researchers involved:Researchers Researchers involved:Researchers Nathalie Gauthier(IRD), Claire Vidal (IRD) Serge Morand(IRD),SylvieManguin(IRD), «Systèmescanniers»(Cirad) (Inra, MontpellierSupAgro, Cirad, IRD) Philippe Ryckewaert, Frédéric Chiroleu, CBGP, Centre deBiologie et Phylogénie-Phylogéographie » Researcher involved:Researcher Populations-Environnement » Serge Kreiter (MontpellierSupAgro), de GestiondesPopulations Researcher involved:Researcher Director: DenisBourguet, [email protected] [email protected] [email protected] Laboratoire Européen Head: Bernard Reynaud, Researchers-teachers involved:Researchers-teachers agricole - Acarologie » Head: Pascal Marnotte, Director: Walker Jones, Unité deRecherche de LutteBiologique USDA-ARS, EBCL, Other teams Jean-Philippe Deguine [email protected] Alain Migeon(Inra) UMR 1062 Bernard Reynaud, SergeQuilici, Denis Bourguet(Inra), Jacques Fargues(Inra), Marie-Claude Bon Régis Goebel

© F. Hérard © F. Hérard • Evolution ofmatingsystems; coevolution ofplantsandinsects; • Evolution ofspecializationand populations; • Evolution oflifecycles instructured evolution. The maintopicsare: natural populations, and experimental demographic andgeneticstudyof development ofstatisticaltools, biology. Methods includemodeling, with anemphasisonconservation populations dynamicsandgenetics control butdevelopsandusestoolsof 1 2 1. Contact: Franck Hérard, [email protected] deciduous trees, makes themaseriousthreat to biodiversity intheseecosystems. nurseries, orchards, andnearby forests. The widerangeofhostplantsthesepests, particularly areas where they are initially introduced are agri-ecosystems, themajorchallengestopreserving Early detectionofthe host plantsiswiderthanthatof hedges (e.g. and grown. Very quickly, many speciesofornamentalswere affected aswell asbushesand this specieshave beenfound attackingmaples neargreenhouses where bonsaiswere stored hand, introductions of East from untreated boards cut upfrom trunksofinfested deciduoustrees. Ontheother of wood packingmaterial, pallets, crates, cablespools, andwedging materialmadeintheFar in urbanareas. Introductions of In otherrespects, thestructuralweakening oftrees, withhighriskofbranchfall, isproblematic heartwood. Attacked trees are unsuitablefor lumberprocessing. feeding inthecambiumandphloemlayers. canbore longgalleriesintothe Olderlarvae visible roots ontheground. tunnelunderthebarkcreating galleries,The early stagelarvae while attacksby beetles establishedinEurope Anoplophora spp.,

Anoplophora glabripennis Anoplophora glabripennis 2.

Anoplophora chinensis Prunus laurocerasus accidentally introduced introduced intoEurope A. chinensis in Europe in2000 Anoplophora by bonsaïtrade A. chinensis are found inthelower oftrunks, portion nearthecollar, andon ), andeven oldrose bushes. The known rangeof spp. infestations andtheeradication ofthesepestsintheurban A. glabripennis A. glabripennis. were madethrough ofbonsais. theimportation Adults of A. glabripennis healthy trees, oronly lightly stressed trees. Attacks by of themajorpests urban areas andinpoplarplantations. damage tothegenera of deciduoustrees. InChina, from theFarEastwhere they live onnumerous species 2000/29/EC and2002/36/EC). Both speciesare originally in theEuropean Community CommissionDirectives (for Europe,eradication efforts theregulations are held emergency measures againsttheirspread, with starting Anoplophora pest toastatusof “invasive species” inthatcountry. with inappropriate control measures, couldraisethis increasing speedofitspopulationgrowth, inconjunction extent ofthecurrent infestation by status of “introduced pests.” InItaly, however, the In Europe atthepresent time, bothpestshave the fruit trees, nurseries, andforests. urban areas. However, bothpests are seriousthreats to on trees andonornamentalshave beenfound only in North America andEurope. Presently, infestations and Two Asian longhornedbeetles, A. chinensis, inthe West occurred through theimportation Assessment of Asian longhorned spp. are quarantinedspeciessubjectto inaequidens and oneinvasive species( corymbosa with tworare especies( The lattertopicsdealparticularly conservation. gene flow, localadaptationand • Fragmentation, demography, are located in the upper portion oftrees are locatedintheupperportion have beenaccidentally introduced into Citrus Populus, Acer, and ). in Japan. Bothpestsattack A. glabripennis

Brassica insularis Anoplophora glabripennis invasive species and A. chinensis A. chinensis Salix Centaurea causesheavy A. chinensis Senecio , principally in andthe isone )

Characterising biodiversity in agroecosystems A multidisciplinary study on a spider mite Tetranychus evansi Baker & Pritchard, a recent arrival in Europe

The spider mite Tetranychus evansi belongs to the Tetranychidae family. This family consists of some 1,200 species worldwide, about a hundred of which are crop pests with about ten of these being pests of major importance. Tetranychus evansi is linked mainly with the Solanaceae and is a threat to tomatoes, potatoes and other related crops. In our region, host plants include Solanaceous weeds that are much appreciated by the mite, providing reservoirs and relay zones. This is the case with black nightshade (Solanum nigrum) a widespread plant and a principal host in Spain. Other plants in the family — including weeds and/ or ruderals — may play the same role in other regions.

Tetranychus evansi appears to have originated in South America where large numbers were observed on tomato crops for the first time in 1954. It was then reported in the United States and later on Mauritius in the Indian Ocean. It was subsequently found throughout the Mascarenes and along the east coast of Africa. © A. Migeon The species was reported in Morocco at the end of the 1980s. A female of the mite Tetranychus evansi At the beginning of the 1990s it was observed in Portugal, in Macaronesia and then in Spain. It was found for the first time in The use of molecular markers makes it possible 1) to differentiate France in the Roussillon region in 2004. between closely related or cryptic species, 2) to trace the history of a biological invasion, and 3) to accurately monitor This history shows that it is a particularly invasive species that recent population developments. The use of genetic markers has succeeded in spreading over a large part of the world in just in spider mite studies has provided a tool for distinguishing half a century. It has caused serious damage to tomato crops between the species. A PCR-RFLP marker has been developed wherever it has been introduced. The damage has been magnified to distinguish between T. evansi and T. urticae, two pests by the cultivation of the crop in new zones. This is the case frequently found on the same crops. The precise study of the in particular for Réunion Island, where the recent increase in dispersal of species in close or similar climatic zones provides tomato production in a protected environment has run up against valuable information. It is thus observed that T. evansi is present the presence of this spider mite. It is also the case in Morocco, throughout Florida. The northern part of the state has relatively Spain and in Roussillon, France, where increasing quantities of cold winters with a fair number of days with frost and average tomatoes are being grown. winter temperatures below 10°C.•••

Before 1970

1970 - 1980

1980 - 1990

1990 - 2000 Biological control Biological After 2000 17

World distribution map and invasion of Tetranychus evansi Migeon A. From 18 Biological control whitefly addition, itshouldbenotedthatlike otherexoticpestssuchasthe Mediterranean zoneandinthefoothills ofthePyrenees. In seems tobemostatrisk. Riskismoderateinthebroader, inland In France, theMediterranean coastalzone, includingCorsica, • average July temperatures 20°Corhigher (thisenablesthe • fewer than40days offrost peryear (conditioninnorthern • average temperatures maximum January of10°C(development • average temperatures January of5°Corhigher(theconditions characteristics already determinedinthelaboratory. These are: determined by its American distributionaswell asbiological Tetranychus evansi ••• technical hurdles, scientistsinBrazilandKenya beganabiological acaricides andthe development ofresistance. Faced withthese the spread of practices norpesticideapplications have succeededincontrolling productioncurtailing ofSolanaceouscrops. Neithercultural implemented, in Kenya inparticular where thepestisseriously Various methodsfor controlling greenhouses throughout Europe. a recent arrivalinEurope Tetranychus evansi A multidisciplinarystudyonaspidermite From A.Migeon months). development ofthree generationsduringthetwo summer Florida); threshold) orhigher; Florida); in northern In order todeterminethepotentialfor thespread of Bemisia tabaci, T. evansi inFrance, we tookintoaccount four criteria populations, tolow efficacyof dueinpart

T. evansi is athreat tocolonizecommercial Baker &Pritchard, T. evansi populationshave been Tixier-Garcin & Serge Kreiter, [email protected] Contacts: Maria Navajas, Alain Migeon, Marie-Stéphane implemented inanoptimum, efficaciousmanner. unintentional introduction willberequired for thesesolutions tobe on theabilitytomassrear them, andonrisksassociatedwith the predators, ontheirabilityto establishinzonesofintroduction, years. Otherwork onthetaxonomicandgeneticcharacterization of potential for effective biologicalcontrol of bioassays.characteristics inlaboratory The results indicatehigh already madeitpossibletofindspeciesexhibitinginteresting T. evansi The populationparametersandthecapacity ofpredation on mites were found, includingspecies thatwere new toscience. three years. Duringthissurvey work, numerous speciesofpredatory This work consistedofprospection indifferent regions ofBrazilfor are alsomites, belongingtothefamily Phytoseiidae. present. The principalandmosteffective predators of Tetranychidae observed, leadingtotheassumptionthateffective predators are T. evansi with collaborationbetween Brazil, FranceandKenya. Indeed, predators inthezonesoforigins results ofthevariouscontrol methods, work onthesearch for around theworld were found tobeineffective. Given thepoor commonly marketed andusedinbiologicalcontrol ofspidermites control program for thispestinthe1990s. However, thepredators iscommonlyinBrazilbutnooutbreaks observed are have beenandstillare beingstudied. This longwork has conditions based onclimatic spread inFrance Tetranychus evansi W of Probability fortheinstallation Forecast of T. evansi Low Negligible Assessment of in France

T. evansi T. evansi High Moderate invasive species was started in2002 wasstarted inthecoming © R. Sforza basin the Mediteranean that isinvasive along from North America a speciesoriginating eleaegnifolium, Solanum

19 Biological control 20 Biological control Researchers involved:Researchers Team «Unitéd’évaluation génétique» Thematic « Bioagresseurs émergents Thematic «Bioagresseurs Mireille Fargette(IRD), Mateille(IRD) Thierry regulations oftheEuropean working onthistheme products, orotherproducts Andy Sheppard (1 Researchers involved:Researchers Directors andresearchers involved: de la diversité descommunautés de ladiversité (Inra, MontpellierSupAgro, Cirad, IRD) de nématodesphytoparasites » import soil,import plants, micro- CBGP, Centre deBiologie et Program « Écologie etgestion de GestiondesPopulations scientific communitycan Mic Julien (2 scientific purposes. These harmful organisms, crop to conditionsconcerning importations areimportations subject onto French for territory Director: DenisBourguet, [email protected] Laboratoire Européen Laboratoire européen [email protected] Director: Walker Jones, Main teams de LutteBiologique and macro-organisms USDA-ARS, EBCL, Other teams [email protected] Any scientistfrom the et bio-invasions » [email protected] Dominique CoutinotandRenéSforza or subjecttospecific UMR 1062 that are prohibited CSIRO, nd semester2006), Patrice Cadet(IRD), st semester2006), Community. agreements, andquarantine issues T Once issued,an agreement isvalid consideration forimportation. under detention ofthe materials management associated with risk applied totheinstallations, and (or institution),maintenance of thepetitioningorganization of theapplicant,personnel quarantine installations, theactivities theapplicable the importation, or objectsunderconsideration in ofthespecies includes descriptions (DRAF/SRPV LR). The fileofapproval ofLanguedoc-RoussillonService Forestry, Regional Plant Protection Regional Director ofAgriculture and outbyThe applicationiscarried the the LetterofAuthority (LOA). accompanied by adocument called under thesestatutesmustbe theterritory entering living material subject topreliminary approval and 22, 2002).Quarantine activitiesare of June 10,1998,decree ofNovember transposed into French law(decree These directives were subsequently in directive 95/44/CEofJuly 26,1995. conditions ofdetention,are specified scientific purposes, aswell asthe in theEuropean Community for introduced orputintocirculation under whichanorganism canbe scientific purposes, theconditions In order toallow for importations prohibited orsubjecttoregulation. products whoseintroduction is the plants, crop products, orother CE ofMay 8,2000,whichalsolists appendices ofdirective 2000/29/ organisms,” appearsinthe also known as “quarantine organisms,A listoftheseharmful and ineffectsinceOctober 2,2005. for foodandagriculture (FAO 1997) the Conference oftheUnited Nations Convention adoptedatthetimeof as includedinthenewtextof for theProtection ofPlants (ICPP) 2.1. oftheInternational Convention or crop products” pathogenic agentharmfultoplants regulatory aspects: or biotypeofplant, animalor defined as he term organism”“harmful is Legislation and according toarticle “any species, strain harmful organisms,harmful also bewater- orairborne. (e.g materials imported soil),orcan whether theorganism isconveyed by containment thatdiffersaccording to present canbesubjecttoalevelof organisms studiedorlikelytobe to theorganisms handled. The accordingcontainment canvary requirements forbiological surrounding community. The as well astheenvironment and protection ofthemanipulators “quarantine” mustensure the The installationofacontainment organization (i.e. DRAF/SRPVLR). the initiativeofadministrative Inspections canbeorganized at to approval mustberespected. which timetheconditionsrelated during for onefive-year period, conform withtheConventionconform on and France. These importations effect withintheEuropean Union the legislationandregulations in species are withregard imported to plants, intotwoquarantines. All and micro-organisms, including onto French livingmacro- territory Control (EBCL)imports Laboratory activities, the European Biological Within theframework ofitsresearch of foreign origin living organisms for thestudyof containment facilities Quarantines and [email protected] and René (USDA-ARS, EBCL) Sforza ** Text by written DominiqueCoutinot [email protected] CILBA AGROPOLIS Co-animateurs duGroupe Informel duConfi ARS, EBCL)andPierre Ehret (DRAF-SRPV) * Text by written DominiqueCoutinot (USDA- [email protected] , [email protected] , né

© D. Coutinot natural enemies, aswell astarget rearing ofpestinsectsandtheir allowing theidentification and containment facilityhasequipment in asingle, contiguousunit. This three greenhouses (each22,5m²) of55m²and up ofalaboratory well assomefrom France, ismade from abroadplants originating as enemies ofpestinsectsandexotic quarantine forthestudyofnatural andplantpathogens. A149m² the identificationandrearing of contains equipmentthatallows level ofcontainment. This laboratory andexitofafacilitythis entry mandated foreachpersonupon showercountries. Anobligatory is pathogens comingfrom foreign the studyofinsectandplant A 25m²quarantine atEBCLallows a LOA. mustbeaccompaniedbymaterials ofregulatedThe importation living (SRPVLR). protection authorities a declaration totheofficialplant organisms are subject to imported are approved andallregulated (CITES). Quarantine activities Species of Wild Flora andFauna International TradeEndangered in with potentialnatural enemiesinplasticbags (e.g. CBGPand CSIRO). International Campus ofBaillarguet and containmentfacilitieson the institutes alsousequarantines control agents, otherresearch pest organisms andtheirbiological Within theframework ofstudieson or invasive plants. biological control againstinsectpests within theframework ofaclassical and/or theirfuture establishment States studies forcomplementary viaaircargo totheUnitedexported these quarantine facilititesmaybe organisms andcultures raised within Under establishedguidelines, personnelonly.is forauthorized Access tothesetwoquarantines each structure. of 60Pascals ismaintainedinside efficiency filter).Anegativepressure filtration system (pre-filters andhigh disposal) andanair-treatment (for processing liquidwastebefore of waste),adecontaminationtank an autoclave(fordecontamination Each quarantine isequippedwith coming from foreign countries. plants andtheirnatural enemies and toolsusedforsecure packaging during travel backtoMontpellier 1. Envelopes forshippingseeds 2. Olives from SouthAfrica 2 1 containment The requirements of • Deliver theapplication totheservice • Assemble application materials for theagreement Different steps • - Appoint anexpert Visit oftheexpert 3.Organisms beingabletobe conveyed 2. organismsabletobe Imported 1. organismsabletobe Imported to levels ofcontainment studied objectsaccording to organismsor Treatments applied • Specifictothe “regulated quarantine • Advice ofthechiefSRPV- Advice • Determinedby ananalysis ofrisk • Prefectoral ordinance • Validation oftheanalysis ofriskby the • Evaluationofthequalitytechnical • Level ofcontainmentaccording to instructor ofDRAF/SRPV - Report and advice of the expert andadviceoftheexpert - Report work organisms” andthetypeofscientific of theDirector oftheDRAF carried outby thepetitioninginstitute expert expert choices andorganization organism(s) the possiblemodeofdispersal containment. be applied tothewhole structure of a positive ornegative pressure can According totheorganismsstudied air: pre-filters andhighefficiency filter. chlorination) andtreatment ofthe of theliquids(hightemperatures, of thesolids(autoclave) andtreatment by thesolids, liquids, andair: treatment (high temperatures, chlorination). (autoclave) andtreatment oftheliquids but notby air: treatment ofthesolids conveyed by solidmaterialsandwater (autoclave). water orair: treatment ofthesolids conveyed by solidmaterialsbutnotby © D. Coutinot 21 Biological control 22 Understanding and evaluating and Understanding Biological control diversity anditsinteractions managers andtechnicaladvisors. research transferlaboratory toland teamswithtechnology land management. These interactions are studiedby field and using suchknowledge tobenefitagriculturalorimprove natural enemies, tounderstandtheirinteractionsand trying Field scientistsare trackingbioinvaders andtheirassociated tropical forests, aswell asMediterranean garrigues anddunes. diversity ofhabitatssurveyed: from central Asian steppesto control challenges. This diversity inapproaches islinked tothe scientists have applied many andvariousdisciplinestobiological following commonecologicalrules. For more thanonecentury, centimeter scale, livingorganismsfrom allecosystemsinteract From thesoiltoair, from themicrometer scaletothe © R. Sforza fullonum) (Dipsacus common teasel seedheads of living in gentianaeana Endothenia lepidopteran ofthe Larva p. 26 genetic diversity Characterization of p. 24 and interactions Evaluating hostspecificity © S. Maurice © R. Sforza research perspectives ontheevolution ofspecialization. its naturalenemies, andtheirco-evolution opensnew genetic structure ofnaturalpopulationsthepest, of for aspecifiedbiocontrol program. Considerationofthe organism were considered, withanaturalenemy selected debated. Before, only behavioural traitsofthebeneficial enemies, inthesearch for host-specificitytraits, hasbeen phylogeographical studiesofbothpestsandtheirnatural ofperforming comparativethe pastdecadeimportance chloroplastic) DNAsequenceinformation. Inaddition, in like microsatellites ororganellar(mitochondrial its geneticvariabilityusingneutralmolecularmarkers and introduced rangesofthetargetpest, measures target species, basedonsamples collected inthenative involved. Inthatregard, phylogeographic analysis ofthe understanding geneticcharacteristicsofthespecies The studyofinterspecificinteractionsisfacilitatedby for naturalenemies. map isobtainedthataidsindirecting foreign exploration presenting similarcharacteristicsworldwide. A climatic invaded rangeofthepestinquestionandidentifiesareas to determinehygrometric andtemperature trends inthe symbiotic organisms). Comparative climateanalysis helps (including plantassociations, parasite complexes, and one tocharacterizetheecosysteminwhichitevolved Pinpointing thecenteroforiginapestspeciesallows design offieldexperiements. complex. Inaddition, studiesofabioticfactorsinform the study interactionsbetween allprotagonists ofabiological evaluate theseinteractions, host-specificitytestingaimsto (e.g. monophagous strainsvsoligophagous). Inorder to predation capabilities orvariabletrophic characteristics different populationsofthesamespecieswithdifferent parasitism andcompetition, orintraspecificamongst These interactionsmay beinterspecificasinpredation, B of variousbiologicalinteractions. field observation, scientificstudy, andmanagement iological control isalongprocess thatbeginswith and Marie-ClaudeBon(EBCL) René Sforza 23 Biological control 24 Biological control Brian Rector, William Meikle, DominiqueCoutinot, accepted thateachinvasive Researchers involved:Researchers Thematic « Bioagresseurs émergents Thematic «Bioagresseurs Researchers involved:Researchers invaded by thepestspecies. themselves. It iscommonly insect, avariety oftestsare Andy Sheppard (1 classical biocontrol agents Directors andresearchers involved: for weedsandinsectpests between anatural enemy range, butnotallofthese to lackofhost-specificity. Mic Julien (2 as biocontrol agentsdue undertaken forselecting undertaken minimum of15natural reasonably host-specific contre les bio-invasions » Team «Luttebiologique In order toevaluate the will notbeappropriate Laboratoire Européen Laboratoire européen An criterion important Janine Vitou, Mireille Jourdan species isattackedby a [email protected] Director: Walker Jones, Main teams de LutteBiologique be released inthearea level ofhostspecificity USDA-ARS, EBCL, and itstargetplantor Evaluating hostspecificity the bestcandidatesto for selectingpotential and notbecomepests [email protected] et bio-invasions » [email protected] enemies initsnative is thatagentswillbe Franck Hérard CSIRO, Walker Jones, RenéSforza, Andy Sheppard, MicJulien, nd semester2006), st semester2006), range. range. protected speciesintheintroduced endangered species, andnative as crops ofhigheconomicimpact, genera andrelated families, aswell in thesamegenus, thenrelated withspeciesandsub-species starting according approach; toacentrifugal 20 to50plantspeciesisselected species. For thatpurpose, alistof a listofcloserelatives ofthetarget species requires testingitagainst a seed-feedinginsectontarget evaluating thehost-specificityof In thecaseofweed biocontrol, organism’s hostrange. thebreadthdetermine ofthe behavioural, and ecologicalfactors Phylogenetic, genetic,physiological, target species by itsnatural enemy. in theselectionandutilizationof invasions team « andinthefield(EBCL, laboratory bioassays are inthe untertaken effects, extensive host-specificity In order tominimize non-target safely employed. biological control organism canbe introduced range, alesshost-specific inthe or economicimportance or exotic speciesofenvironmental no closelyrelated nativespecies in effect,ifaninvasive specieshas upon thetarget speciesselected; The needforspecificitydepends protected andendangered species). non-target organisms (e.g. crops, feeding oroviposition behaviouron non-target effectssuchasaccidental such aspredators), of the lesstherisk (as opposedtogeneralist enemies The more specificanatural enemy, in biologicalcontrol natural enemies Evaluating specificityof Lutte biologiquecontre lesbio- »).Many factorsplayarole and interactions uncontrolled populations than expected,withwidespread outtobemoreturned poplyphagous for controlling cereal aphidsbut Harmonia axyridis predators suchaswhentheladybug, unintended effectsmayoccurwith cannot becomepests. Nevertheless, natural enemiesofweeds, parasitoids In addition,unlikephytophagous degree ofspecificityfortheirhost. and moths, oftenpresent ahigh pests suchasscaleinsects, aphids, crop insects usedagainstimportant are fasterandeasier. Parasitoid for natural enemiesofinsectpests In biocontrol tests comparison, authorizations. of thetarget plant,underlegal release intotheintroduced range of rearing), itmaybepetitionedfor significant impactonthetarget, ease desirable traits (i.e. specificity, If theagent selectedpossesses open-air dependingonthesituation. either inaquarantine facilityorin greenhouses andgrowth chambers, conditions ingarden plots, conducted undercontrolled These multiple-year testsare consideration.them from further suitable hostsandquicklyeliminating which plantspeciesare definitelynot providing aneasywaytoascertain species atatime)are conducted, No-choice tests(i.e. testingoneplant potential hostrange ofaherbivore. are toassessthe bothperformed Feeding activityandoviposition tests plants havebeguntoproduce seeds. oncethe controlled experiments together withthenatural enemyin sources, thengrown andplaced from botanicalgardens andother Seeds ofeachspecies are obtained [email protected] * Text by written René (USDA-ARS,EBCL), Sforza wasintroduced * . pathogen biologicalcontrol strategy the developmentofanefficient Mycoparasite/Antagonist systemfor Study ofinteractions inaPathogen/ mycoparasitic andantagonistic properties andefficientcontrol ofthepathogen andMichel Ducamp, [email protected] species of (i) evaluation ofthediversity andpopulationdynamicsof designed adevelopmental project withbasicscientificquestions involving biologicalcontrol and: Institut deRecherche pourledéveloppement Agricole underlying theinteractionsbetween pathogen underthepressure ofthismycoparasite aswell asresearch intothemechanisms this decisionrequires studyofthepopulationdynamics, genetics, andadaptive capacity ofthe would beagood biologicalcontrol agentinthefieldanddeveloping methods tohelpreach bioassaysin laboratory undercontrolled conditions. Determiningwhetherthiscandidate Several isolatesofthefungus control ofBPD. consideration hasbeengiven toanintegratedpestmanagementapproach, includingbiological fungicides andalsobecompatiblewiththeculturalpracticesof African farmers, more need todevelop strategiesthatwillbothprotect theenvironment through minimaluseof Contacts: Marie-Claude Bon, [email protected] Pathogen/Mycoparasite/Antagonist complex. functional genomics); and(iii)studyofthemechanismsunderlying the interactionswithinthe Cocoa pods of apathogenicfungusonwidevarietyplantspecies S Settingupagarden experimenttoevaluatespecificity Phytophthora in Africa; (ii)investigation oftheir evolution intimeandspace(through Given theabsenceofdisease-resistant varietiesofcocoaandthe Trichoderma asperellum How tooptimizethecontrol in Africa, andinCameroon afflicting upto90%oflocalproduction. Phytophthora intensification ofdiseasepressure, mainly duetoBPD, in particular, ofthemostaggressive casualagentofthe

plantations. iscurrentlyThis strategy constrained by the © M.C. Bon Black Pod Disease( is concentrated, istoincrease thesizeofcocoa cocoa? How tominimizethecostoftreatments and in Africa, where 69%oftheworldwide production to preserve theenvironment whilemaintainingthe demand for cocoa, thegeneraltrend amongfarmers In order tosatisfytheincrease inworldwide quality ofcocoa? These are new challengesfor the biological control ofBPD. and (IRAD)inCameroon, EBCLandCiradhave (host-specificity test) collectedinCameroon have shown Trichoderma Trichoderma asperellum Phytophthora megakarya, © R. Sforza . Incollaborationwiththe P. megakarya andthemain BPD)of

Ustilago phrygica caput-medusae of thegrass T Philippe Ryckewaert, FrédéricChiroleu, Jean-Philippe Deguine Laboratoire Européen deLutteBiologique Marie-Stéphane Tixier-Garcin (Montpellier SupAgro) naturels, Écoledes Mines d’Alès(EMA) bioagresseurs enmilieutropical, Cirad/ Laboratoire Géniedel’environnement Experimental contamination Experimental industriel etdesrisquesindustriels Researchers involved:Researchers Team «Unitéd’évaluation génétique» Team leader: PVBMT (Peuplements végétauxet Team «Écologie animaleetZoologie working onthistheme des cultures pérennes »(Cirad) « Systèmescanniers»(Cirad) (Inra, MontpellierSupAgro, Cirad, IRD) CBGP, Centre deBiologie et « Maîtrisedesbioagresseurs Serge Kreiter (MontpellierSupAgro), Unité Propre deRecherche Researcher involved:Researcher Director: MiguelLopez-Ferber, de GestiondesPopulations [email protected] Researcher involved:Researcher Université delaRéunion) Researcher involved:Researcher Director: Bernard Reynaud, Director: DominiqueBerry Director: Pascal Marnotte, [email protected] [email protected] Director: DenisBourguet, [email protected] Researchers-teachers involved:Researchers-teachers agricole - Acarologie » Director: Walker Jones, Unité deRecherche USDA-ARS, EBCL, Christian Cilas, [email protected] Other teams Taeniatherum [email protected] UMR 1062 UMR C53 bythesmut Bernard Reynaud, SergeQuilici, LGEI, Marie-Claude Bon Christian Cilas Régis Goebel

© R. Sforza 25 Biological control 26 Biological control technology have spawned technology the modesofcolonization techniques formeasuring genetic variation thatare now available toexplore of foreign speciesinnew such asagriculture and consequences onworld agriculture. Theboost global trade increases native ranges. Rapid species outsidetheir in humanactivities geographical areas. molecular biology molecular biology have undesirable improvements in the movement of a broad range of Bioinvasions Characterization of ARS) focuson: Agricultural Research (USDA- Service States ofAgriculture Department (EBCL) oftheUnitedLaboratory the European Biological Control Genetic Evaluation of Laboratory The mainresearch activities ofthe Lepidium draba each protagonist ( of history also oftheevolutionary context ofecologicalconstraints but (e.g. hostrange) notonlywithinthe analysis ofdifferential interactions is mutuallyinformative, allowing phylogeography foreachprotagonist specific andthecomparative plant communitiescanbehighly Host-herbivore relationships in history.its evolutionary markers, allows thereconstruction of its natural range usingmolecular which canbeassessedthroughout The geneticdiversityofaspecies, are openingnewperspectives. being takenintoconsideration and evolution ofspecialization,are now are instudyingthe instrumental weed and itsnatural enemies, which natural populations ofboththetarget architecture andlocalevolutionof thegenetic Data concerning biological control context. insects were being chosenforagiven species were takenintoaccountwhen species andcombinationsofsuch behavioral traits ofnatural enemy now, onlythestudyofcertain biological control ofweeds. Until specialization incandidatesfor enemies insearch oftraits forhost- of hostplantsandtheirnatural the comparative phylogeography ofstudying community onthemerits past decadewithinthescientific A debatehastakenplaceinthe plant the target natural enemyfor Selecting theright genetic diversity p.29 see box related to ). (see box, p.29) by the weevil of theinvasive weed Rome (Italy) thecontrol concerning (Montana, USA)andUniversity of inSidneythe USDA-ARSlaboratory biological control ofBPD;andwith (UK) regarding theproject onthe and Imperial College ofLondon Développement de laRecherche Agricolepourle level includingwiththe initiated attheinternational Several collaborations havebeen regions. for control ofBPDincocoagrowing to exclusive reliance onfungicides approachto propose analternative SupAgro/Cirad). The ultimategoalis Plant Protection (Inra/Montpellier plants parasites forIntegrated and Geneticofinteractions between phytopathogènes “ withCiradin partnership (team These research activitiesare led agent in response toabiologicalcontrol ( of Black Pod Disease ofcocoa thepathogenicity) (in particular • The studyoftheadaptivecapacity SupAgro andCBGP. in collaboration withMontpellier species. This approach isconducted effective managementoftheinvasive specific genotypes)withthegoalof (i.e. subspecies, orpopulation- the mostspecificentitiespossible populations andtoassignthem geographical ofinvading origin main objectivesare to trace the and theirnatural enemies. The species (weeds orinsectpests) phylogeography ofbothinvasive • The analysisofcomparative [email protected] * Text by written Marie-Claude Bon (USDA- ARS, EBCL), Phytophthora megakarya Biologie et génétique des eucaryotes Biologie etgénétiquedeseucaryotes (Trichoderma asperellum) Ceutorhynchus assimilis * . ••• (IRAD)inCameroon ”, theUMR “Biology Lepidium draba Institut , BPD) .

© D. Navia and spread ofexoticpests Using geneticdiversitytotrace theorigin observed genetic variationissummarizedonthe nextpage.observed respectively, were amplifiedby PCRandsequenced. The 16S region. These two fragmentsof1000and404basepairs, of theribosomalclusterandafragment ofthemitochondrial regions: thenuclear internaltranscribed region (ITS1-5.8S-ITS2) region). DNAsequencevariation wasexaminedintwo genomic been reported(the Americas, Africa, andtheIndianOcean most ofthegeographic areas where thespecieshascurrently relationships of29populations elucidate theputative source ofthepest, we analyzed thegenetic In order toinvestigate the invasion process ofthespeciesandto of origincoconut. not beenrecorded intheIndo-Pacificregion, thepresumed area concern IndiaandSriLankainthe1990s. Curiously, themitehas most recent records ofthespread ofthemitetonew areas in 1967, andlaterinEast Africa, in Tanzania inthe1980s. The in West Africa, intheGulfofGuineaislands1966, andinBenin already beenpresent inSouth America. The mitewasreported in 1965. However, priorrecords certain indicatethatithad unknown. The specieswasfirstdescribedinGuerrero, Mexico, of theproduction areas ofthatcrop. The originofthemiteis pestofcoconut andhasrecentlyan important spread tomost mite, pest,economically important thecoconut mite. This eriophyid traitswasusedtoinvestigatehistory theinvasion pathways ofan A combinationofgeneticmarkers andinformation onlife- populations andthespread oftargetedpests. This information canbeusedtodeterminetheoriginsof organisms through bothhistoricalandevolutionary timescales. markers are idealfor tracingthemovement patternsof Because ofthenature ofgeneticmaterial, neutralgenetic Aceria guerreronisAceria , hasemergedover thepast30years as A. guerreronis originatingfrom 1 © D. Navia © D. Navia ••• locus 3 locus 2 locus 1 A. Phylogeneticapproach B. Assignmentbasedapproach haplotype studied potential originofpopulations Genetic approach topinpoint the originof populations potential originofpopulations 3 guerreronis mite phytophagous by the caused in coconut Outbreaks 1. 2.3. Aceria 2 haplotype studied

27 From M. Navajas Biological control 28 Biological control Following theoriginalmitochondrialhaplotypes America andanothersinglehaplotypewas haplotype wasfoundinCentral andNorth the nuclearITSsequencesrevealed thatall (a) The variousmitochondrialhaplotypes non-American samples(inred) show very (b) Congruently, thetree with constructed are represented inseveral branches ofthe highest nucleotidediversity wasfoundin whereas theBrazilian samples(ingreen) dots) are indicatedbythepiecharts. The types were present. Bycontrast, onlyone in thedifferent sampledlocalities(black (in blue)are gathered inasinglecluster. 1 haplotype shared byallnon-Americanmitesfrom Brazil where sixoutoftheseven haplo- tree. The rest oftheNew World samples detected andtheirrelative proportions Samples ofeachsamplingregion are Phylogeographical history Africa andtheIndian subcontinent. little diversity andclustertogether, colour-coded asonthemap. of thecoconutmite. 1 haplotype 6 haplotypes Contact: Maria Navajas, [email protected] represents aclearandimpending threat. other Asian andPacificcountrieswhere ithasnotyet beenreported butwhere it using quarantinemeasures inorder toprevent thespread ofthisdestructive miteto ofany palmtreetransport propagation materialthusrepresents arisktobemanaged from ortradeofhost-plantpropagation humanactivityinthetransport material. The It iswidely acceptedthatmostspread ofthecoconut mitetoremote areas results long distances. germplasm exchange)have clearly facilitatedtherapid disseminationofthepestover of mitesover distances, short humanactivities(e.g. internationaltrade, tourism, and a realistic hypothesis. Although wind isconsidered tobethemain meansofspread of themitewithin Americas andfrom there toothercontinentsappears tobe unique infestation oftheseregions, presumably from the Americas. The dissemination polymorphism between African andIndo-Pacificsamplesiscompatiblewitharecent and Some insightsintothespread ofthepestare provided here. The absenceof a distinctgenotype. mites collectedonanotherpalmanalyzed here the miteoriginally fed onapalmtree otherthanthecoconut palm. evidence,As further question oftheoriginalhostplantmite. apreviousThis supports hypothesis that One issueraisedby thehypothesis ofaNew World originof there. mite inthe Americas ishigh, we sampledtheentire hadonly partially diversity ofspecies continents’ samples. Oneexplanation couldbethatbecausegeneticvariationofthe none oftheNew World haplotypes identifiedinthisstudywere present intheother With thehypothesis ofaSouth American originofthemite, itremains unclearwhy most likely occurred inSouth America. diversity isfound inthe Western Hemisphere andsuggests thatthisoriginofspecies the practically simultaneous reportofthemitein Americas and Africa, thehighest throughout theworld ofthispest. helptotracetherecent expansionhistory Despite ••• and spread ofexoticpests Using geneticdiversitytotrace theorigin Molecular results togetherwithinformation onrecords ofthepresence ofthemite 1 haplotype Characterization of (Syagrus romanzoffiana) Following ITSnuclearsequences A. guerreronis genetic diversity , donotrepresent isthe From M. Navajas Americas Indo-Australasia region and Africa Ceutorhynchus How toselecttherightnatural enemy The casestudyof Contacts: Marie-Claude Bon, [email protected] program targetingthisweed. this information willbetaken intoaccountintheplanningofon-going biologicalcontrol suggest recent andmultiple introductions of genetic diversity andthemultiple geographical originsofthe US populationsoftheweed oftherecent colonization of history by thispolyploid species.American territory The high of two intergenicregions ofchloroplast DNAfrom theweed hashelpedinretracing the historical andspatialfactorsonthehost-specificityofthisnatural enemy. Molecularanalysis of thetwo host-races. This studyshouldallow scientiststodeterminetheinfluenceofboth andJean-François Martin, [email protected] weed unlikely. program for thisweed considered theexistenceofahighly specificbiologicalagentagainstthis is cruciferous like cabbageandrape. Recently, researchers involved inthebiological control Lepidium draba innovation isbasedonthedirect has beendevelopedatEBCL. This gall weevil, non-invasive, ofthe whitetopcollar- A methodoftyping,bothdirect and destroying them? signature ofsuchspecimens without it possibletoobtainthegenetic endangered species. How then,is a program ofan ofconservation control program againstapestor hinder thesuccessofabiological differentiation ofgenotypes andcan present significantchallenges to morphological phenotypescan entities, identical despitevirtually of distinctgeneticandbiological presence withinoneinsectspecies candidates, orpestspecies. The endangered insects, biologicalcontrol applications aimedatstudying interestingdoor tosomeparticularly molecular techniquesopensthe The developmentofnon-invasive the fieldofentomology assay ofgreat value to A non-invasive genetic assimilis Understanding andevaluating ssp. Ceutorhynchus assimilis showed acongruencybetween thegeneticandgeographic distributions draba However, differences inbehavior andspecificitywere between observed similar profile oftheweed initsnative area. The comparisonclearly

© M. de Freitas , aweed thatistoxic tocattle andisinvading American rangelands, populations ofthewhitetopcrown-gall weevil, phylogeographical profile oftheweevil compared was further toa reared from different hostplants. Sequenceanalysis ofmitochondrial the existenceofaracenaturalenemy specifictotheweed. The barrier betweenraceandtheothersconfirmed thisparticular DNA regions from many different populationsidentifiedseveral The observation ofhostspecialization andofareproductiveThe observation distinct genotypeswithinthespecies, allmorphologically similar one racethatwashighly specificto but differing inhostrangeandgeographic distribution, including Lepidium draba , Lepidium draba diversity anditsinteractions information thatwasnotpossible information from insect frass isproviding new first studybasedonDNAextracted candidates towards theirhosts. This on specificityofbiologicalcontrol essential intheframework ofresearch can behighlighted--factorsthatare of analysis, somebiologicalfactors they belong.From thesetwolevels as well asthehostrace towhich identification ofindividualinsects Molecular markers allow sequences.differing stranded DNAfragments with slightly electrophoretic ofdouble- properties technique measures differences in Conformation Polymorphism). This technique DSCP(DoubleStrand region, combinedwiththe markers from agivenmitochondrial is basedonpolymorphismin the insecthasleftplant.Detection insectspecies,a particular evenafter also allows onetodetectfeedingby typing ofDNAobtainedfrom frass the fecalexcretions oftheinsect. The genotypesdirectlyof cryptic from genome, allowing thedifferentiation detection ofregions oftheinsect’s ssp. Lepidium draba draba Ceutorhynchus assimilis Ceutorhynchus from Eurasia. All of : ssp. draba . , ragwort). Senecio inaequidens guerreronis different models(seeboxes): two teamsare interested intwo of exotic bio-aggressors. These characterize thegeneticdiversity using molecularbiologytoolsto Sciences del’Évolution) et environnement bio-agresseurs Processus évolutifs etgestiondes environnement gestion dessystèmespopulations- The teams “ genetic ofinvasions Diversity ofthemodels in quarantine operations. research anddevelopmentincluding could beappliedtootherareas of destroyed. In thefuture, thismethod approach. No insectsare disturbedor alone orthrough aclassicalgenetic to getthrough fieldobservations

Team «Écologie intégrative desSystèmes Team «Unitéd’évaluation génétique» Team « GénétiqueetEnvironnement » Institut desSciencesdel’Évolution (Inra, MontpellierSupAgro, Cirad, IRD) Researchers involved:Researchers CBGP, Centre deBiologie et Researcher involved:Researcher Team «Génétiqueadaptative Populations-Environnement » de GestiondesPopulations Researcher involved:Researcher [email protected] Researcher involved:Researcher Director: DenisBourguet, [email protected] Laboratoire Européen Director: NicolePasteur, Director: Walker Jones, Main teams de LutteBiologique USDA-ARS, EBCL, (thecoconutmite)and (Montpellier SupAgro) [email protected] Écologie intégrative et Alain Migeon(Inra) et spéciation» (UM2, CNRS) ) and “ UMR 1062 ” (CBGP, Team ” Jean-François Martin Maria Navajas (Inra), Sandrine Maurice Marie-Claude Bon (Institut des (narrow-leaved Génétique are also Aceria 29 Biological control 30 Biological control Philippe Ryckewaert, Chiroleu, Frédéric Jean-PhilippeDeguine Michel Martinez (Inra),Michel Martinez Jean-François Germain(LNPV), Philippe Reynaud (LNPV), Jean-Claude Streito (LNPV) Marie-Stéphane Tixier-Garcin (MontpellierSupAgro), Marie-Stéphane Tixier-Garcin (Montpellier SupAgro),(Montpellier Marie-Stéphane Tixier-Garcin (Inra), BrunoMichel(Cirad), FranckDorkfeld (Inra), naturels, ÉcoledesMinesd’Alès(EMA) bioagresseurs enmilieutropical, Cirad/ Laboratoire Géniedel’environnement Researchers involved:Researchers Brian Rector, William Meikle, DominiqueCoutinot, Naturelle, assignedby IRD), Armelle Cœur-d’Acier industriel etdesrisquesindustriels UM1, UM2, UM3, MontpellierSupAgro, Cirad) Researchers involved:Researchers Bernar Johan Michaux(FNRS)GauthierDobigny (IRD), Gérard Delvare (Cirad), Mireille Fargette(IRD), Thematic «Systématique-Taxonomie, Laurent Granjon(MuseumNationald’Histoire Researchers involved:Researchers PVBMT (Peuplements végétauxet Team «Écologie animaleetZoologie Jean-Pierre Quéré(Inra), MariaNavajas (Inra), Thematic « Bioagresseurs émergents Thematic «Bioagresseurs working onthistheme [email protected] Researchers involved:Researchers Director: Jean-Dominique Lebreton, Serge Morand(IRD),SylvieManguin(IRD), Andy Sheppard (1 (Inra, MontpellierSupAgro, Cirad, IRD) Director andresearchers involved: CBGP, Centre deBiologie et Researcher involved:Researcher MiguelLopez-Ferber Phylogénie-Phylogéographie » Serge Kreiter (MontpellierSupAgro), Director: MiguelLopez-Ferber, de GestiondesPopulations Mic Julien (2 John Thompson, Anne Charpentier John Thompson,Anne [email protected] Researcher involved:Researcher Université delaRéunion) fonctionnelle etévolutive CEFE, Centre d’écologie Director: Bernard Reynaud, contre les bio-invasions » Director: Denis Bourguet, Team «Luttebiologique [email protected] [email protected] Laboratoire Européen Laboratoire européen [email protected] Researchers-teachers involved:Researchers-teachers agricole - Acarologie » Director: Walker Jones, de LutteBiologique USDA-ARS, EBCL, Other teams [email protected] et bio-invasions » [email protected] (UMR 5175, CNRS, Alain Migeon(Inra) Franck Hérard UMR 1062 UMR C53 Arnaud Martin, Cindy Adolphe, CSIRO, LGEI, Walker Jones, RenéSforza, nd semester2006), Denis Bourguet(Inra), st semester2006), Mireille Jourdan d Reynaud, Serge Quilici, • Contact: SandrineMaurice, [email protected] to invade theEuropean climate. wool from thetetraploid area ofthenative rangeand/orthatonly tetraploids are able by diploids. This suggeststhatthedifferent European countrieshave their allimported Only tetraploids have invaded Europe whileSouth America and Australia are invaded from allopolyploidization (hybridization ofdiploidswithdifferent genomes). the intersectionofthesetwo areas. This patternsuggeststhattetraploids are issued in southern Africa withdistinctgeographical distributions. The tetraploids are found in hasrevealed• Flow cytometry two distinctdiploidgenomes(differing inDNAquantity Polyploidization hasthusprobably occurred insouthern Africa. ofintroduction inEuropethe history suggestsseveral independentintroductions. European • All Lesotho. • This speciescomplexincludesdiploidandtetraploid forms in South Africa and biological speciesbutrepresent acomplexofinterbreeding subspecies. and African originsof agents oneneedstoknow theoriginsofinvading individuals. Studyofthetaxonomy In order tostudytheevolution ofabiologicalinvasion ortofind biologicalcontrol invasion different asitcolonizesbioclimaticzonesvery from thoseofitsnative range. institutes for understandingthebiogeography, ecology, andevolution ofaplant’s pastures throughout theworld andhasbecomeamodelsystemfor various Agropolis As such, animal-dispersed seeds. months ofgerminationandindividualscanproduce upto10,000long-lived, wind-and threatens livestock andnative plants. Itsextensive flowering withinafew period starts Mediterranean climatesupto2500minaltitudewithabroad ecologicaltoleranceand Introduced asawool contaminantatMazametin1936, itgrows well intemperateto and pastures, itisnative tocoastalriver systemsandstony pastures ofsouthern Africa. A tetraploid, toxic, herbaceous, tussock-forming perennial found invineyards, dunes Europe andisthedominantinvasive plantspeciesintheLanguedoc-Roussillonregion. invasive alienplantspecies, ispredominant among5suchspeciesthroughout Western Senecio inaequidens in Languedoc-Roussillonregion Narrow-leaved ragwort Senecio inaequidens, Senecioharveianus S. inaequidens S. inaequidens DC. (Asteraceae), amemberofgenus thatiswell known amongst Senecio inaequidens is arepresentative ofagroup ofrelated speciesthatinvade plantsthathave beenstudiedare tetraploid, though Characterization of and hasleadtothefollowing conclusions: Senecio madagascariensis From S. Maurice genetic diversity are notdistinct southern Africa southern native range in ragwort inits of narrow-leaved tetraploid forms of diploidand Cartography Flowers of Senecio inaequidens (Asteraceae): one the 5 most noxious weed species in Western Europe

© S. Maurice

Colony of the Towards new associations to aphid, Aphis jacobeae, control narrow-leaved ragwort heavily attacking While the potential of classical biological control has not yet been fully explored, very narrow-leaved high specificity in any agents would be required given the number of native Senecio spp. in ragwort in Europe. Solution-oriented research in Europe focuses on two natural enemies: a rust fungus the vicinity of of Australian origin, Puccinia lagenophorae Cooke, that is now cosmopolitan and can kill Montpellier susceptible genotypes in Europe (though resistance is present in many populations); and the ragwort aphid, Aphis jacobaeae Schrank, which heavily infests the flowering shoots, severely reducing seed set. These agents are being explored for their potential in inundative or inoculative biological control. Biological control Biological Contact: Andy Sheppard, [email protected] 31

© A. Sheppard Colonies of the scale insect Neopulvinaria innumerabilis © R. Sforza on grapevine Controlling populations

Within the concept of sustainable agriculture, controlling pest populations through non-chemical approaches is the final step of a long process. It encompasses study and analysis of invasive and emerging species and their interactions with other species and abiotic factors in the habitat they colonize. ‘Controlling’ does not imply complete annihilation of target populations, but establishment of a social, environmental, and economic threshold below which practioners, agronomists, and growers estimate acceptable losses. Biological control and chemical control can be integrated to reduce pest populations below that threshold. Bioloical control Bioloical

32 p. 44 sustainable agriculture another visionof Agroecology: p. 40 for sustainableagriculture Sustainable plantprotection p. 34 introduced natural enemies control: theuseof Classical biological © J.P. Deguine © T.Mateille © F. Hérard sustainability incrop production. agrochemical products, which isoneofthemainideals spurred legislationcallingfor thereduction intheuseof to thepublicopinionandenvironmental awareness that These different managementstrategiesare connected considered. role ofreservoir zonesfor beneficialfaunaispositively preserve native andconserve naturalenemiesandthe systems ornaturalenvironments. The ideaisalsoto management ofplantpopulations, whetherincropping aims toimprove animalandplantbiodiversity through degraded landscapes. Inthisperspective, agroecology and environmental hazards; andtorestore ecologically large quantitiesofhigh-qualityfood; toprevent health social andenvironmental stakes: tosafely produce Agroecosystems are now thetargetsofnew economic, plant anditsassociatedfauna. micro-organisms, thatcoexistandinteractwiththecrop conditions, soilstructure, andspeciescomplexes, including based onconsiderationofnaturalelementssuchasclimatic of environmental interactionsaimsatpestmanagement safety andenvironmental quality. Inthiscontext, thestudy Key componentsofsustainablecrop production include onlegumes,importance cereals, andtrees. sustainable managementtoolsagainstpestsofeconomic or ingreenhouses, suchnaturalenemiesare working as against insectormitepests. Releasedinopenhabitats parasitoids, entomopathogenicfungiandbaculoviruses Thus, moststudiesinEurope concernwithusingpredators, development asnoenemy hasyet beenreleased of biologicalcontrol early ofplantpestsisatavery stage States, Australia, New Zealand, andSouth Africa, thestudy and monitored long-term. InEurope, unlike intheUnited plant management. These introductions are controlled introducing naturalenemiesand, lessfrequently, invasive of invertebrate pestssuchasinsectsandmitesby Classical biocontrol at Agropolis addresses management interactions andagroecology. T classical biologicalcontrol, environmental include are independentandcomplementary hree sustainablepestmanagement practicesthat and Jean-Philippe Deguine(Cirad) René Sforza (EBCL) in natura . 33 Biological control 34 Biological control requires amultidisciplinary through natural regulatory native area ofthepest. This Classical biologicalcontrol Classical biologicalcontrol: mechanisms ofpredatory utilizing natural enemies exotic oremergingpests reduce andlimitexotic is a strategy tomanage is astrategy populations. It aimsto originating from the pest populationsby scientific approach. the useofintroduced natural enemies (CBGP) becauseofagood systèmes population-environnement “ bug)” waschosenby theteam Macrolophus caliginosus tabaci Bemisia greenhouse tomato(crop plant) / system The tritrophic “Mediterranean lagged behindthatofotherregions. research activityinthisarea has the 1980s. However, European systems hasbeenestablishedsince population dynamicsincomplex of populationbiology. Analysisof with clearlinkstothediscipline populations (pestsandantagonists) involves in-depthstudiesoftarget management. Such anapproach biocontrol-based integrated pest be prominent componentsof and safetyconsiderations should making implementation.Quality by datafordecision- experimental strategies and tacticssupported requires thedevelopmentof Pest managementinagroecosystems greenhouse tomato Bemisia tabaci- Macrolophus caliginosus- tritrophic system: Dynamics ofthe Écologie intégrative etgestiondes (major pest)/ (predatory ” by abaculovirus. to thenatural control ofthisinsect Languedoc (France) are due inpart Lymantria dispar example, thecyclic of variation host populationdensity. For epizootics are in linkedtovariations in fieldhostpopulations. Such frequently causenatural epizootics several years). Baculoviruses time between hostencounters(upto allows of forlongperiods itssurvival system, theocclusionbodythat have developedaprotection of plantsorinthesoil. These viruses environment ontheleavesandfruits Baculoviruses canbefoundinthe baculoviruses Insect pestcontrol by environment, etc.). agroecosystem (i.e. cropping, physical with theotherkeyelementsof integration of the IPMcomponent biodemographic mechanismsand to developabetterunderstandingof essential tools. Clearlythere isaneed management whereas itsIPMlacks the crops withthehighestlevelof in thatgreenhouse tomatoisoneof agroecology. There isagreat paradox understanding ofthedynamicsits populationsinthe •••

on a (Hymenoptera: Trichogrammatidae)moving 2. stalks onReunion Island 1. The eggparasitoid Stem borer infestingsugarcane larvae Chilo saccahriphagus the caseofaugmentativereleases of the sugarcanestemborer onReunionIsland: Towards biologicalcontrol of was toolow for effective pestcontrol. Trichogramma chilonis parasitoids showed thepresence ofonly onespeciesonRéunion, biological control programs worlwide. A fieldinventory ofthese known andeffective eggparasitoidsthatare commonly usedin spp. (Hymenoptera: Trichogrammatidae), agroup ofwell- of parasitoidstocontrol theborer, amongthem Cirad andFDGDON. Onthisisland, there isanexistingcomplex control program onRéunion, in closecollaborationwithInra, Council of a project co-fundedby theEuropean UnionandtheGeneral interesting andenvironmentally friendly solution. Over 3years, insects are usually insidethestalk, biologicalcontrol presents an are difficulttoapply andcanbeineffective sincethetargeted more susceptibletotheborer. While chemicaltreatments compared to100120t/hafor theothervarieties)thatare yield sugarcane varieties(suchas varietyR579; yieldof150t/ha this phenomenonislinked tothedevelopment ofnew high- reported onMauritiusandReunionislands. OnReunionisland, and sucrose quality). Recentstemborer outbreaks have been indirect lossesduringprocessing (reduced recoverable sugar (upto40t/ha)and terms ofcanetonnagepriortoharvesting caused by itslarvae. Suchfeeding causesdirect lossesin for significantcrop lossesduetointernaldamageinthestalk Chilo sacchariphagus countries bordering theIndianOcean. The spottedstemborer, strongly impedeproductivity, insouthern particularly Africa and countries. However, stemboringinsectsattackingthiscrop Sugarcane crop isaneconomically important inmany tropical La Réunion Trichogramma chilonis (Lepidoptera: Crambidae), isresponsible , andthenaturaldensityofthisparasitoid allowed theestablishmentofabiological eggbatch

Trichogramma Trichogramma 1423 production by 15to36tonsofcane perhectare (i.e. +571to releases) thaninthecontrol plots. This increased sugarcane internodes inthetreated plots(thosewith (2002-2004) ledtosignificantly lower percentages ofbored production andfieldexperiments. Three years offieldreleases control agent. This strainwastherefore chosenfor mass climate) appeared tooffer thebestpotentialasabiological A populationoriginatingfrom Saint-Benoît(hotandhumid chilonis control andmassrearing, thebiologicalcharacteristicsofthree To choosethewaspsoffering thebestabilitiesfor bothbiological corresponding tothemostactive oviposition periodoftheborer. T. chilonis Our objectiveoutaugmentative wastocarry releases of select more effective T. chilonis process, torefine itisnowthe releaseof necessary strategy To ensure profitability andreliability ofthisbiologicalcontrol biological control using allowed optimisticaboutthegeneralizeduseof ustobevery hot andhumidareas ofLaRéunion. Suchresults from smallplots density wasthehighest, asituationusually associatedwiththe The bestresults were obtainedinthe plotswhere thehost Contact: Régis Goebel, should leadtofuture successesintheseendeavours. and development thatprevailed partnership duringthisproject € strainswere compared inthelaboratory. /ha). inthecanefieldsatbeginningofcrop cycle, (periodsanddoses), inaddition tocontinuing to Trichogramma Trichogramma [email protected] © D. Conlong 1 populations. The research by thesurgarcane farmers. Trichogramma © R. Goebel T. 2 35 Biological control 36 Biological control naturels, ÉcoledesMinesd’Alès(EMA) bioagresseurs enmilieutropical, Cirad/ Laboratoire Géniedel’environnement Researchers involved:Researchers in additiontodifferent modesof diseases andweeds inagroecosystems innovative control methodsforpests, These research activitiesshouldleadto islands. terrestrial ecosystemsinremote endemism andinvasion ofnatural resistance topestsanddiseases; genetics andcharacterisation ofplant interactions,dynamics andtritrophic populations); pestpopulation of phytopathogenicmicro-organism andadaptation epidemiology (survey La Réunion.Its fieldsofstudyare plant centre atSaint-Pierre, inthesouthof ofthecrop protectionunit ispart with University ofLaRéunion. This terrestrial ecosystems, inpartnership of crops andbiodiversityconservation focused onprotection oftropical bioagresseurs enmilieutropical PVBMT, in theJoint Research C53 Unit (JRU The research andtraining activities conducted onLaReunion Research activities industriel etdesrisquesindustriels PVBMT (Peuplements végétauxet « Systèmescanniers»(Cirad) Philippe Ryckewaert, Frédéric Chiroleu, Director: MiguelLopez-Ferber, [email protected] Université delaRéunion) Researcher involved:Researcher Director: Bernard Reynaud, [email protected] [email protected] Head: Pascal Marnotte, Main teams Unité deRecherche Peuplements végétauxet Jean-Philippe Deguine UMR C53 Bernard Reynaud, SergeQuilici, LGEI, Régis Goebel plant diseasevector Bemisia tabaci, both apestand in greenhouses whitefly thatis a Hemipteran ) are Contacts: OlivierBonato, [email protected] IPM decision-makingtools. crop-based, field-testedmodelto provide information leadingtothe development of them inmulti-species pestmodels andfinally toimplementthemintoanintegrated approach. The mainchallengeistodevelop mechanisticsubmodels, toincorporate toshiftfromimportance anempirical approach toamore rigorous, predictive the dynamicsofmultispecific systemsbasedonadequatemodelling isofprimary (iii) simulation modelingofthedynamicsmultitrophic system. The analysis of to temperature andhostplant(e.g. development models, life-table analyses), and parameters of bothpestandnaturalenemy(ii) life-history populationsinrelation of populationsover spaceandtime(e.g. samplingplans, populationstructures), The French program isfocusedaspects: onthree complementary (i)distribution developed for greenhouses equippedwithinsect-proof screens. focicontrol ofthewhitefly invasion theprimary basedonaprophylactic IPMstrategy (Roussillon andProvence), aFrench collaborative research program wasinitiatedto andJacques Fargues, [email protected] mainly basedonLaRéunion,is whose scientificandtechnicalstaffis on sugarcane systems(Cirad), farming resistance.varietal The research unit gical control, agronomic practices and are puttingmore emphasisonbiolo- In thiscontext,research institutions (e.g. stalkborers). developmentofsomepests internal high densityofthecanestalksand because ofthelongcrop cycle, the insecticide treatment on sugarcane to othercrops, there islittleorno plantdiseases).Asopposed borne leaves (e.g. thrips, aphid- armyworms, stalks (e.g. mealybugs, stalkborers) or affects eitherroots (e.g. whitegrubs), real economicincidence. The damage insects, ofwhich10specieshavea The sugarcane crop attracts numerous against sugarcane pests Biological control organisms. areas andmanagement ofinvasive forindegenousforestconservation © J. Fargues and economiccontextofthetargetedproduction regions approach hasbeenplanned. Taking intoaccountthesocial to thishighly complexproblem, IPM amultidisciplinary at several sitesin2003. Sincethere isnodirect solution and the Tomato Yellow LeafCurl Virus (TYLCV)wasdetected invaded thevegetable production areas ofsouthernFrance Recently, the whiteflyisadoublethreat Pest andvector: Bemisia tabaci in France. Bureau ofSugar Station Experiment Research Institute (MSIRI),andInra (SASRI), theMauritius Sugar Industry Sugarcane Research Institute developed withtheSouth African plete. To has thisend,apartnership to sugarcane growth remain incom- the pests. Population modelsrelative fauna thatinteract withandregulate fields tobetterexploitthearthropod study ofbiodiversityinsugarcane There isaneedformore detailed blishment ofinvasive pests. mate ontheemergence ortheesta- impact ofagronomic pratices andcli- willcontinuetoinvestigatethe effort chilonis sacchariphagus nic fungusandthestemborer marginalis control ofthewhitegrub ned onLaReunion in thebiological Outstanding results havebeenobtai- ment amongothersubjects. interested inintegrated pestmanage- Classical biologicalcontrol biotypes, vectors ofBegomovirus,

(seebox). The research by anentomopathoge- with Trichogramma Hoplochelus Chilo moth larvae on moth larvae cabbage leaves diamondback Anoplophora glabripennis Host/parasitoid interactions ininvasive • tobetterknow • todetermine A. glabripennis spread.further Studiesofhost/parasitoidinteractions appropriate methodtoavoid outbreaks ofthesepestsandtheir the pestsestablishinforests, biologicalcontrol couldbethemost Moreover, iferadicationofinfestations inurbanhabitatsfails, orif ofthepests. enemies attackingeggsandearly stagelarvae the otherhand, couldbereached effectively by specificnatural Infested trees thatmightbemissed by themonitoringofficers, on some attacked trees are notlocatedduringinitialinspections. that are oftentiny ormasked. Therefore itisinevitable that inventory theattacked trees isavisualdetectionofthesymptoms of theinfestations buttheonly techniquecurrently available to The control monitoring ofthesepestsisbasedonpreliminary eradication fails. to theireradicationand(ii)abatethepestpopulationsif problem ofthe pests? Biologicalcontrol canbehelpfulattwo levels tosolve the Why andhow tousebiologicalcontrol toattackthesetwo (Cotesia plutellae) Study ofdiamondbackmothanditsnatural enemy East; predators andevaluation oftheirimpactonthehostsin Far enemies intheirnative area -inventory ofparasitoidsandthe pests ashostsin known European parasitoids from thelocalfaunahave already acceptedtheexotic new associations); thefirststep consistsindeterminingifsome biological control agentsagainstthesepests(i.e. evaluation of of taxonomy, host-plants, andbehaviour would beeffective cerambycids thatare closely related to Damage by Controlling Diamondback mothadult (Lepidoptera: Plutellidae) and Anoplophora ifsomenaturalenemiesofEuropean A. chinensis thecomplexof populations © D. Bordat spp. introductions: (i)tocontribute are considered intwo ways: Anoplophora Anoplophora populations nativetodifferent areas

Anoplophora © D. Bordat Contacts: DominiqueBordat, [email protected] • Moleculardifferentiation: by RFLPofpolydnavirus DNA • Geneticdifferentiation: detectionofgeneflow between populations(DBMandnaturalenemies) • Allozyme differentiation ofpopulations(DBMandnaturalenemies) • Biologicaldifferentiation: comparisonofoviposition behavior, (DBMand morphometry Four parallelstudieswere carried outtocharacterizethishost/parasitoidcomplex: divergent behaviors are geneticinorigin. three speciesdiffered noticeably. Inorder tosetupIPMprograms toknow ifthe itisimportant undernaturalconditionsindifferentour observations areas, thebehavior ofpopulationsthese Probably native totemperatezones, bothare alsooftencollectedintropical climates. During (Hymenoptera : Eulophidae)are effective naturalenemiesofDBMandare specifictoDBMlarvae. Cotesia plutellae (e.g. 20 generations/year intropical areas. Excessive useofchemicalpesticides, includingbio-pesticides six developed continents. pestintemperateclimates,A secondary DBMhasmore than pests ofBrassicaceaecrops (e.g. cabbage, radish, turnip, rapeseed) worldwide andispresent onall Plutella xylostella andLaurence Arvanitakis, [email protected] mitochondrial DNA(COI)andby ISSRmethod(DBM). C. plutellae spp. natural spp. infestations. B. thuringiensis spp. interms ), parasitim%, life-cycle time, sexratio(naturalenemies) (Kurdjumov) (Hymenoptera : Braconidae)and (L.)(diamondbackmoth, DBM)(Lepidoptera: Yponomeutidae) isamongthemain and ), insomeregions hasledtotheappaerance ofresistant strains. and Gérard Delvare, [email protected] Contacts: Franck Hérard, [email protected] the localecosystems). (following appropriate testsontheirimpactnon-targethosts in parasitoids couldbeusedinbiological control by augmentation If eradicationoftheseexoticpests fails, itissupposedthatthese Italy todiscover otherpotentialantagonists of is inprogress atEBCLandexplorations continue inthefield A. glabripennis identified, andfour speciesamongthemalsoaccepted From thisstudy, sixnew associations with were already known inEurope from otherxylophagous hosts. (L1andL2)of larvae Anoplophora spp. These parasitoids Sclerodermus Westwood (Hym.: , Pteromalinae), and Pteromalidae, ), (Hym.: Eupelmidae), (Hym.: Eurytomidae), Braconidae), Spathius erythrocephalus to from theFarEastwithitshost)isspecific likelyEulophidae) (whichwasvery introduced accidentally parasitoid infested with of thepestsatseveral siteswithinandoutsidethearea by exposingsentinelplantsinfested withvariousstages these pests. InItaly, parasitismofbothhostswasstudied and evaluate European parasitoidsthatcouldhelpcontrol biological control studieswere implemented toidentify brochure. Inconjunctionwiththeeradicationprograms, of thepromising parasitoidspeciesare shown inthis Some progress hasbeenmadeinthesetwo fields; afew A. chinensis A. chinensis. Aprostocetus anoplophorae Eurytoma melanoneura Eurytoma sp. (Hym.: Bethylidae) attacked early stage asahost. The evaluation ofsomethesespecies A. chinensis. Six larval ectoparasitoids, Sixlarval brevis Calosota vernalis Wesmael (Hym.: Itwasshown thattheoophagous (C. plutellae), Trigonoderus princeps populations Oomyzus sokolowskii Walker Delvare (Hymenoptera: Boucek(Hym.: Curtis Curtis by sequencingof A. chinensis Anoplophora

(Kurdjumov) were were spp. 37 Biological control

© F. Hérard 38 Biological control Marie-Stéphane Tixier-Garcin (Montpellier SupAgro) (Montpellier Marie-Stéphane Tixier-Garcin « Productions fruitières» ethorticoles Team «Écologie intégrative desSystèmes Brian Rector, William Meikle, DominiqueCoutinot, Brian Rector, William Meikle, DominiqueCoutinot, Researchers involved:Researchers involved:Researchers Team «Écologie animaleetZoologie Olivier Bonato(IRD), FrédéricPellegrin (IRD), Thematic « Bioagresseurs émergents Thematic «Bioagresseurs Mireille Fargette(IRD), Mateille(IRD) Thierry working onthistheme UPR Horticulture, Département Nathalie Gauthier(IRD), Claire Vidal (IRD) Researchers involved:Researchers Andy Sheppard (1 Researchers involved:Researchers Directors andresearchers involved: de la diversité descommunautés de ladiversité (Inra, MontpellierSupAgro, Cirad, IRD) Researchers involved:Researchers de nématodesphytoparasites » CBGP, Centre deBiologie et Populations-Environnement » Program «Écologie etgestion Serge Kreiter (MontpellierSupAgro), de GestiondesPopulations Mic Julien (2 contre les bio-invasions » contre les bio-invasions » Director: Denis Bourguet, Team «Luttebiologique Team «Luttebiologique [email protected] Laboratoire Européen Laboratoire Européen Laboratoire européen [email protected] [email protected] Researchers-teachers involved:Researchers-teachers agricole - Acarologie » Director: Walker Jones, Director: Walker Jones, Head: Philippe Vernier, de LutteBiologique de LutteBiologique USDA-ARS, EBCL, USDA-ARS, EBCL, Other teams [email protected] et bio-invasions » [email protected] [email protected] Laurence Arvanitakis Franck Hérard Franck Hérard UMR 1062 (Flhor, Cirad) CSIRO, Walker Jones, RenéSforza, Walker Jones, RenéSforza, nd Lymantria dispar semester2006), Jacques Fargues(Inra), Patrice Cadet(IRD), st Dominique Bordat, semester2006), Lymantria dispar 1. Caterpillarof Quercus ilext 2. Adultof on © F. Hérard throughout theMontpellierarea Lymantria dispar 1 lepidopteran specieswithoutdistinction. an invasion by thepest; however, ithasalocalnegative effect by killingmany non-target proportion ofL2). The “Bt” treatment canalsoprotect surrounding urbanareas against (withacertain when thepopulationsofpest contain amajorityoffirstinstarlarvae Bacillus thuringiensis occur over several consecutive years inhighvalueoakwoodlots, aerialtreatments witha mothnaturalenemiescanusually controlof gypsy thepesteffectively. When defoliations invertebrate andvertebratepredators. In Western Europe, itisrecognized thatthecomplex mothnucleopolyhedrosisdue tothegypsy virus(NPV), andtoattacksby parasitoidsand many larvae. are survivors Starved weakened andgenerallysusceptibletoepizootics very complete development ofalltheindividuals. occurs, Starvation resulting inthedeathsof this case, thecaterpillars’need for food exceeds thecapacity ofthehabitattoensure the infested forests can betotally defoliated over hundreds oreven thousandshectares. In Gypsy mothpopulationsdevelop incyclicaloutbreaks. Duringtheculminationphase, the duetoskincontactwithyoungexperienced urtications larvae. forests around Montpellier. Inaddition totheenvironmental damage, somelocalresidents many locally intown, ornamentals wasobserved aswell asthroughout largeareas ofoak to parksandprivateyards withinthecity. Duringthefollowing weeks, heavy defoliation of oak forests andwest locatednorth ofMontpellier, were millionsoffirstinstarlarvae borne gusts ofwindthatoccurred motheggswere duringtheperiodwhengypsy hatchinginthe anditincreasesbirth thesizeofinfested area considerably. In2005, becauseofstrong This phenomenontendstolimittherisksofoverpopulationatsite andstarvation threads produced glands. by theirsalivary They canbeblown onthewindfor longdistances. population density, thetiny, disseminateby newly hatchedlarvae “ballooning” onlongsilk and are commonon Gypsy moth Contact: Franck Hérard, [email protected] (Lymantria dispar) (Lymantria (Bt)basedpreparation may bewarranted. The treatment must beapplied Quercus pubescens caterpillarsare defoliators ofoakandmany otherhardwoods and , amothwellknown Q. ilex © F. Hérard 2 intheMontpellierarea. high Undervery Controlling populations

Insect pest control using baculoviruses

The host range of each baculovirus is different and they are formation. Other gene products interfere with host physiology or often quite specific. For example, the Spodoptera exigua multiple behaviour (e.g. climbing before death). Ongoing research aims to nucleopolyhedrovirus (SeMNPV) is extremely pathogenic to better understand baculovirus host specificity and to characterize Spodoptera exigua larvae, an agricultural pest originally from Asia their life cycles and the modifications they provoke in the infected and now cosmopolitan. However, this virus cannot kill larvae from host. The number of completely sequenced baculovirus genomes is other very closely related species, including other Spodoptera spp. increasing, allowing a comparative genomics approach. Conversely, the Spodoptera littoralis nucleopolyhedrovirus (SpliNPV), isolated from S. littoralis, a species found around the Mediterranean, Within a single virus species phylogeographic variation can be can also kill S. frugiperda, a new world species, as well as S. exigua. The found. Baculoviruses adapt to their host populations and these mechanisms involved in baculovirus host specificity have not yet been host populations are not strictly similar. Virus isolates from a given fully characterised. They are not limited to entry into the larval cell, location are often more active against their native host population but correspond to a series of blocking points at various levels of the than against foreign populations of the same host species. Moreover, larval invasion (tissue barriers) and to the replication cycle inside the within a single viral isolate (obtained from a single location at a given insect cells. time) there can be genetic variation. Virus genomes are not uniform and differences in biological activity might be found between them. Baculoviruses are used to control the larvae of certain insects that In some situations, this variability is required for the optimal activity cause losses in agriculture and forestry. Some baculovirus species of the whole isolate, with the various virus genotypes playing are also pathogenic to mosquitoes and might be useful for their complementary roles. control as well (e.g. Culex nigripalpus nucleopolyhedrovirus). The main advantages of baculoviruses are: their host specificity, which explains For a given location, the genetic composition of the viral isolate may the absence of side effects against non-target insects; the possibility change over time. This could result from co-evolution between the of using the same application equipment for baculoviruses as for pathogen and its host. It has been shown that baculoviruses have a chemical insecticides, and relatively easy storage of those biological long history of co-evolution with their respective hosts, although it is insecticides. Insecticidal preparations containing baculoviruses are theoretically possible that a baculovirus could colonize a new host. commercialized for use against insect pests in greenhouses (e.g. to control Spodoptera exigua on tomatoes, peppers, etc.) in storage Contact: Miguel Lopez-Ferber, [email protected] houses (e.g. to control the potato tuber moth, Phthorimaea operculella), © D. Ray and M. López-Ferber in orchards (e.g. for protection of apples against the apple moth, Cydia pomonella), in row crops (e.g. for protection of soybeans against the Occlusion bodies velvelbean cartepillar, Anticarsia gemmatalis) or in forestry (e.g. for control of Lymantria dispar in hardwood forests). of baculoviruses induce natural epizootics in insect Baculoviruses enter the larva per os, when it feeds on contaminated pest populations leaf, flower, or fruit surfaces. One of the limits for baculovirus efficacy is the need for the presence of the virus where the larvae are feeding. The borers (like the apple borer or the potato tuber moth) enter inside the fruit or the tubers soon after hatching, and once inside, they cannot be treated. This behaviour necessitates that the application of the virus should be prior to boring. For leaf- feeding caterpillars, contamination can occur at any time during larval feeding. Baculovirus virions are protected from degradation by a protein layer that coalesces into a crystalline form. This stable structure is called a polyhedron or granule due to its form, and more generally, an occlusion body. Occlusion bodies typically dissolve in the larval midgut. Occlusion body production is not an exclusive characteristic of baculoviruses: virions for two other virus families infecting arthropods also produce occlusion bodies. However, occlusion body production has never been observed outside the -infecting viruses.

The larva dies due to massive viral replication. By the end of the infection cycle, the larva is more or less a bag of occlusion bodies. When the integument breaks, the occlusion bodies are liberated into the environment, ready to resume the infection cycle. Often, infected larvae are induced by the virus to climb upwards on the host and die in a characteristic position, hanging from the undersides of leaves, which facilitates the spread of subsequently liberated occlusion bodies.

The virus life cycle is complex. Baculovirus genomes are between 80 and 150 kb in size and can encode 100 to 150 genes. A fraction of these genes encodes the structural proteins involved in virus particle control Biological 39 40 Biological control fruit andvegetable production. (Integrated Pest Management) the sametimeanaimcapable agricultural chemicalsisakey Sustainable plantprotection goals ofthereduction by 50% of attracting popularinterest United States proved tobeat factor. Therecent, unfulfilled agriculture, particularly in environmental qualityare of theuseconventional The decrease intheuseof of 75%thecrops inthe pesticides inEurope orof at thecore ofsustainable the integrated protection Pest managementand and autopiandream. or sustainableagriculture i. The segmentofresearch aimed thelast30years: during Production. So itcanbeshown that agents inIntegrated Biological agents andofthemicrobial antagonistic and assessmentofbiologicalcontrol R&D programs, ofintegration interms approaches thatare implementedin knowledge andtherelevance ofthe enables ustoquestionthestateofour An analysisofrecently publishedworks economic communities. technicalandsocio- the concerned withthewholeof in partnership projects developed multidisciplinary the emergence ofanemphasison and sustainabledevelopment,with facing innovation incrop protection on thestakesandstumblingblocks few years, adebatehasbeenopened reach, from a few cases. apart For a constraints, haveremained outof socio-cultural and economic level four, wideningittoconsider tactical setofthecrop systemand three, integrating thestrategic and strategies ofcrop protection. Level level withtheintegration ofallthe of cases, theyhavereached asecond number given crops and,inacertain regulate majorpestpopulationson methodsinorderof alternative to through afirstlevelofdevelopment development programs havegone Conceptually, mostresearch and sustainable development Farm productions and at aconstantlevel ; remained relatively low andnearly integrated production systemshas as biologicalcontrol agentsin at theuseofmicro-organisms of circumstances. which canbeappliedtoawide range assessment, anddecision-support, relying onmethodsofsimulation, a crop systemshouldbespecifiedby control agentsintomanagementof conditions ofintegration ofbiological of theproduction system. The considered asoneofthecomponents and theirnatural enemiesshouldbe and populationdynamicsofpests other constraints). Thus, pestcontrol agroecosystem (crop, economic,and antagonists) andtheconstraints ofthe predators, parasitoids, microbial populations (i.e. invasive species, interactions between protagonistic to explainthedynamicsof the needforasystematicapproach organizations haveclearlyrealized Having this, mostresearch observed vi. The gapbetween reductionist v. Research inecologyand iv. So-called «academic»research has iii. Specialization by discipline, and ii. Reductionist approaches have (IRD), [email protected] * Text by written Jacques Fargues (Inra) andOlivier Bonato disciplines hasincreased. disciplines andintegrative of progress ;and epidemiology havenotmadealot level ; of mechanismsatthepopulation level attheexpenseofanalysis of mechanismsatthemolecular systematically favoured theanalysis divisions amongresearchers ; by scientifictheme, hasreinforced integrative approaches ; developed attheexpenseof * . ••• , [email protected] roots ofmelon nematode infesting Meloidgyne ii. To developandtestglobal i. To setuptheaconceptualbasisfor are bothacademicandapplied: them. describe The aimsofthegroup mechanistic mathematicalmodelsto systems anddevelopmentofessentially eco-epidemiological processes ofthese group hasbeentheinvestigationofkey The scientificapproach favoured by this Bemisia »). approche systémiquedurisque méditerranéens ettropicaux : gestion derisquesenagrosystèmes populations-environnements et « (Theme in agricultural environments studies complexpopulationsystems Montpellier SupAgro, Cirad andIRD) « CBGP»(UMR1062–Inra, A group ofresearchers from economic disciplines scientific andsocio- ofthe at theinterface A systematicapproach strategies ofthis typeof species ;and invasivearthropods, particularly linkedtooutbreaksrisks ofpest methods tomanagephytosanitary the developmentofsustainable Dynamique desystèmes Dynamique spp. rapid responses tosuchthreats, rely onefficientinspectionsand linkedtobio-invasions will risks control of thephytosanitary Underwarming. theseconditions, and travel aswell asglobalclimate the intensificationofglobaltrade common inthefuture becauseof is expectedtobecomemore of aninvasive pest. This phenomenon the introduction andestablishment created crisis bymajor phytosanitary approach toa for amultidisciplinary (see box) exceptionally dangerous plantvirus d’Azur region an andvectoring and intheProvence-Alpes-Côte now proliferating inRoussillon thatis species oftropical origin whitefly Current research iscentered onthe to invasive species. linked risks emerging phytosanitary implemented inorder tomeet organization policiestobe centered onregulation and socio-economic disciplines, scientific (e.g. biophysical)and of thinking, attheinterface group hasworked onsystematic management. Since 2005,this . It isanemblematicmodel Bemisia tabaci Bemisia , aninvasive group’s research activities. enable theimplementationofthis ofthe«CBGP» in NSIperimeters) growth chambers, andphytotrons (molecular biology, greenhouses, facilities experimental to thevarious entomology andmycology andaccess The equipmentofthelaboratories of de laProtection des Végétaux, etc. des Chambres d’Agriculture, services et Légumes, Stations Expérimentales Technique Interprofessionel desFruits ( and institutionalpartners as collaboration withprofessional conception vsfeasibility)aswell at eachstageoftheproject (i.e. l’action etledéveloppement (Inra « /Department of Maine) and integrative disciplines Agroalimentarias, de Investigacion yTechnologia IRD, Inra, Cirad, IRTA- include collaborations withCNRS, academic disciplines(whichcurrently on strong interactions between This research program relies prediction tools. effective diagnosis, simulation,and necessitating thedevelopmentof and theUniversity Sciences pour El Instituto Centre ») © T.Mateille ). 41 Biological control 42 Biological control bioagresseurs enmilieutropical, Cirad/ « Productions fruitières» ethorticoles Team «Écologie intégrative desSystèmes Researchers involved:Researchers PVBMT (Peuplements végétaux et Olivier Bonato(IRD), FrédéricPellegrin (IRD), Mireille Fargette(IRD), Mateille(IRD) Thierry working onthistheme UPR Horticulture, Département Researchers involved:Researchers Nathalie Gauthier(IRD), Claire Vidal (IRD) de la diversité descommunautés de ladiversité Researchers involved:Researchers (Inra, MontpellierSupAgro, Cirad, IRD) (Inra, MontpellierSupAgro, Cirad, IRD) Researchers involved:Researchers de nématodesphytoparasites » Philippe Ryckewaert, FrédéricChiroleu, CBGP, Centre deBiologie et CBGP, Centre deBiologie et Populations-Environnement » Program «Écologie etgestion de GestiondesPopulations de GestiondesPopulations Université delaRéunion) Director: Bernard Reynaud, Director: DenisBourguet, Director: DenisBourguet, [email protected] [email protected] [email protected] [email protected] Main teams Head: Philippe Vernier, Other teams Jean-Philippe Deguine Laurence Arvanitakis UMR 1062 UMR 1062 (Flhor, Cirad) UMR C53 © J. Fargues Bernard Reynaud, SergeQuilici, a boundary-layerstory Climatic constraints tofungipathogenicwhiteflies: Jacques Fargues(Inra), Patrice Cadet(IRD), Dominique Bordat, Contacts: JacquesFargues, [email protected] system. view ofthegreenhouse ventilation, canopy transpirationactivity, andcrop-growth estimate theimplicationsfor microbial control ofwhitefliesfrom thepointof humidity insidethegreenhouse andinleafboundary-layer wasinvestigated to to thesurrounding airingreenhouses. The effects ofgreenhouse ventilation on considerably withinadistanceof5mmfrom theundersideofleafwithrespect laminar flow wasdeveloped andtested. Itwasshown thatairhumidityincreased A modelofmoisture transfer occurring withintheleafboundary-layer under spite ofsignificantdifferences inventilation rates. with vent openingsurfacesrangingfrom 7to19%), thefunguswasnotaffected in ventilation systems(sophisticatedglasshousevs. polyethylene-covered greenhouse, was nodifference infungusefficacy. When comparingtheinfluenceof greenhouse spite ofsignificantdifferences inairflows (0.7-1.2and0.3ms-1, respectively) there of nymphs locatedonlateral-row plantstothatofnymphs oncenter-row plants. In heterogeneity wastaken intoaccountby comparingthefungus-inducedmortality “humid” compartment, aswell as inthe compartment.“dry” The climatic bytreatments >85%inboththe reducedwhitefly larvae numbers ofsurviving thaninthe compartment one.“dry” Inspiteofthisdifferential, mycoinsecticide period ofhighhumidity(>90%RH)wastwo tothree timeslongerinthe “humid” by openingtheridgevents 2hlessthanusualduringthenight. Thus, thedaily greenhouse climatewasfirstmanipulatedfor optimizingmycoinsecticide efficacy efficacy inprotected tomatocrops. Becauseofexpectedclimaticconstraints, the to estimatetheimpactofMediterranean climateconstraintsonmycoinsecticidal Eight small-scale, replicated greenhouse trialswere performed insouthernFrance remainedfungus-induced mortality athighlevels. Consequently, underambientconditions expectedtobeunfavorable (RH<86%), humidity intheaircirculating intheimmediateenvironment oftheleafsurface. measurements demonstratedthattheleaves significantly increased therelative cells underaconstantly circulating flow ofhumidity-regulated air. Microclimatic Bioassays were carried outonintactleaves sandwichedbetweenplastic airtight andClaire Vidal, [email protected] Sustainable plantprotection humidity onatomato Microcaptors ofair leaf surface in Mediterranean tomatogreenhouses. microbial control ofthesemajorpests nymphs toassessthepotentialfor hyphomycetous infection ofwhitefly on theclimaticdeterminismof Roussillon region, hasfocused by theLanguedoc- supported a collaborative research program, on environmental factors. Therefore, are considered tobetoodependent widely underestimated sincethey control agentsispromising butstill development asinundative biological The potentialofthesefungifor tabaci plant suckinginsects, e.g. are abletoregulate populationsof of infection, entomopathogenicfungi Because oftheirpercutaneous mode forsustainableagriculture in California and Texas. Bemisia of bananatree infesting aroot Radopholus similis and thesoilenvironment Nematode-parasitic bacteria The bacterium are notwell known. interactions between nematodepredators andtherhizosphere taking littleornoaccountoftheimpactenvironment. Thus, of biocontrol experimentswhile agentsismeasured inlaboratory soil environments andtohost-specificityissues. Acclimatization by exoticbiocontrol agentstounfamiliarclimaticconditionsor Mediterranean regions, remains deficientduetoweak adaptation economicthreshold,up toasatisfactory especially intropical and capacity tocontrol thedevelopment ofplant-parasiticnematodes (e.g. greenhouse cultures, high-valuecrops). Moreover, the pursued onlyspecificagronomic invery andeconomicsituations plant-protection, nematodebiocontrol ispoorly developed, being Compared tootherresearch anddevelopment areas regarding P. penetrans contributes tothevariabilityofefficacy obtainedwhenusing areas, orindividualplots. The heterogeneity ofsoilfactors distribution scale; whetherworldwide, regional, across production islands. However, variable, itsabundanceisvery regardless of the in sub-Arctic regions, inthetropics andsubtropics, andon cosmopolitan, having beendetectedinalltemperateregions, the level ofbothprey andpredator populations. the absenceofitshost), adensity-dependentbalancecontrols is anobligateparasiteofnematodes(i.e. unabletoreproduce in populations by preventing theirreproduction. As thisbacterium are thefavorite targetsof 90 different genera. Root-knotnematodes( more than200speciesofnematodesbelongingtomore than for nematodebiocontrol. Pasteuria penetrans

P. penetrans hasbeenreportedfrom whichreduces nematode Meloidogyne P. penetrans spp.) is Contact: Mateille,Thierry [email protected] integrated managementrelevant andefficient. manures, rotations, resistant plantvarieties), make meso-biological systems andstrategies(e.g. irrigation, tillageregimes, organic knowledge oftheenvironment andproper managementofcrop frequency ofvariousindigenousbiocontrol agents. Improved Management ofsoilfactorscouldincrease theefficacyand repulsion between thetwo organisms. ionic bridges. The soilsolutionmay alsodecrease hydrophobic concentration ofthesoilsolution, whosecationswould actas surface glycoproteins oftheorganismsaccording totheionic Spore adhesioncouldbedeterminedby thesaturationof the electrochemical fieldsofbothnematodesandbacteria. binding protein interactions. Attachment can beinfluencedby cuticle andabiochemicalrecognition ofthecuticlethrough distinct steps: arandombiophysical connectiontothenematode The attachmentofthespores tonematodesproceeds intwo infest nematodes. colloids inclay soils. Leachedspores andadsorbedspores cannot quickly leachedinsandysoilswhilethey are strongly adsorbedon P. penetrans clay andsandy-clay soils. of The studyofthetransport high proportions ofinfected nematodesare usually detectedin Although nematodepopulationsizesare oftenhighinlightsoils, infesting aroot of banana tree penetrans Pasteuria spores insoilcolumnsshowed thatthespores are © T.Mateille © T.Mateille 43 Biological control 44 Biological control with nematodesandmicro- also affectthecommunity example, thesoilinteracts preventing pestoutbreaks conservation biology. The of organismsthrough the taking intoaccountlarge This reasoning integrates management ofhabitats characteristics ofthesoil scales ofspaceandtime. and pestscommunities, organisms andtheflow or eradicating them, by rather thancontrolling structure over time. For physical andchemical including landscape challenge consistsof multiple disciplines The agroecological management, and ecology, landscape sustainable agriculture substrate. as formore traditional systems, for intensiveagriculture aswell The approach isthusrecommended production. preserves theobjectiveofdurable last recourse. But agroecology also isthevery management; chemistry forward theecologicalcharacter of includes different processes putting The agroecological approach as onthescaleoflandscape. fauna isusefulonalocalscaleaswell forbeneficial biological corridors control. For example, therole of pests, whichcanenhancebiological populations ofnatural enemiesof The process alsoaimstopreserve size; arrangement; composition). structurenot cultivated (form, and the plantsettlements, cultivated or an emphasisonmanagementof and plantbiodiversityby placing management aimstoenhanceanimal not satisfactory. Agroecological paradigm ofcrop protection is of thesituation:chemical Everyone realizes theambiguity compounds. treatments basedontoxic active consequences ofphytosanitary reduction are secondary important the trophic webs, andbiodiversity pollution oftheenvironment and Acute andchronic intoxications, activities andotherlandscapeuses. prevalent unsustainableagricultural to • topropose socialalternatives landscapes; • torestore ecologicallydegraded environmental hazards; • toprevent health and sufficient quantities; • tosafelyproduce qualityfoodin stakes: A social andenvironmental objects ofneweconomic, groecosystems are now the another visionof Agroecology: Agroecology: of the risks of pollution,etc)*. of therisks safeguarding biodiversity, reduction (respectcriteria oftheenvironment, decisions), andenvironmental social (shared andaccepted (traditional tolerance thresholds), account atthesametimeofeconomic outlook,taking and withalong-term thresholds, onthelandscapescale recognition ofnew(andevolving) commons, landscapes, etc.); and on aglobalscale(e.g. watersheds, analyzed tofacilitatedecision-making obtaining localdatathatcanbe economics, etc.) ;scalemodification, agronomy, crop protection, socio- interdisciplinarity (includingecology, the scientificlevel:integration and major methodologicalstakesat invasive specieselucidatesthree settlements andpopulationsof Lastly, managementofplant return toanecologicalbalance. toa biodiversity andtocontribute fauna oninvasive species, torestore theactionofbeneficial to support functions. Associationsofplantstend more specieswithcomplementary specieswithoneor one harvested speciesorof of several harvested temporal orspatial associations multi-species systems, basedon notes astrong renewed interest for the useofagrichemicals. One dead endinthecontextofreducing monoculture isnow regarded asa Except forsomenotable exceptions, plantspecies.of complementary is theuseoffavorable associations useful formanagementofcrop pests One agroecological techniquethatis agriculture.industrial and unintentionalconsequencesof knowledge andavoidthenegative while seekingtodeveloplocal [email protected] * Text by written Jean-Philippe Deguine (Cirad), ••• prophylactic vs.ecologicalstrategies Pest managementforgreenhouse tomatoes: The introduction of vaporariorum andBemisiatabaci. and several mycoinsecticides tocontrol thewhiteflies Trialeurodes (e.g parasitoids,(e.g predators, andentomopathogens). provide better-adapted guidelinesfor selectionofantagonists natural enemiesused by inoculative release must be studiedto Consequences ofthesemicroclimatic changesontheefficacyof confinement, withdaily increases in temperature andairhumidity. France). Screen-equipped vent openingscreated conditionsof were detected(TYLCVisadeclared quarantineorganismin pulling upofthewholecrop assoonthefirst TYLC symptoms a well-adapted solutiontotheFrench regulation whichrequires the for decreasing theriskofwhitefly adultintroduction appeared tobe and continuous heatingduringcoldperiods). Insect-proof screens cultivars insophisticatedgreenhouses year-round (virtually culture is mainly achieved by long-cyclecropping ofvirus-susceptible tolerance thresholds. Tomato production for thefresh-fruit market a new approach totheplantprotection becauseoflow strategy Yellow LeafCurl Virus (TYLCV)intosouthernFrancenecessitates formosa,Encarsia Eretmocerus eremicus, bug such asthepredatory The European tomatoIPMprogram involves 10naturalenemies the greenhouse biocontrol market. largest. These three companiessupply more than75%of Europe has26natural-enemy producers includingtheworld’s three crop ofthistrend remains the greenhouse-grown tomato. Today inEurope. important has beenparticularly The mostemblematic integrated pestmanagementprograms inallmainvegetable crops During thepasttwo decades, theprogression ofbiocontrol-based B. tabaci Macrolophus caliginosus populationstransmittingthe Tomato and Eretmocerus mundus ; theparasitoid ; Contacts: Jacques Fargues, [email protected] sustainable tomatoproduction inMediterranean Europe. approaches canbedeveloped toovercome the “ model). According tothesedifferent contexts, two mainpestcontrol enemies oftargetedpestsfrom theoutdoors(e.g. Catalanproduction closedstructures,partially allowing naturalre-colonization by natural largely isolatedfrom theoutside(e.g. production inRoussillon)and are two groups ofMediterranean greenhouses; sealedenclosures Depending onthegeoclimatic, social, andeconomiccontexts, there enemies (augmentationbiologicalcontrol). severe pestoutbreaks require releases ofmass-produced natural control) cancontrol whitefly populationsinmany situationsbut antagonists biological intoandoutofthegreenhouses (conservation biological control. The continuous migrationofthesenatural andaugmentativeas thedominantspecies)for bothconservation and Olivier Bonato, [email protected] favorable to tomato crops are cultivatedinunsophisticatedtunnels. Though to only pullupthoseplantsshowing TYLC symptoms. Short-cycle In contrast, inCataluña, theSpanishregulation requires farmers risk causedby region, iscommittedtoasystematicapproach tomanagementofthe The research program, by theLanguedoc-Roussillon supported E. mundus focused onthepotentialoflocalnaturalenemiessuchas natural control. An ecologically basedapproach ofIPMhasbeen wide tomaximizeintroduction ofthesepopulations, whichcanexert develop inareas neighboringtomatogreenhouses. Vents are opened of tomatoproduction inMarocco 1. Experimental greenhouse Experimental and a guild of predatory mirids(including andaguildofpredatory 2. B. tabaci, Hygro-thermic sensor in thetomatofoliage B. tabaci themildclimateenablesnaturalenemiesto of agreenhouse anditsphytovirus inthisprophylactic context. © Th. Mateille © Th. Bemisia risk M. caliginosus ” for

© J. Fargues 2 45 Biological control 1 46 Biological control on theevolutionoftheirlifetraits (phylogeny,history phylogeography) affect them,toevaluate theimpactof environmental interactions that communities andonbiological order toacquire knowledge on nematode modelisrelevant in on agroecology. The plant-parasitic or landscapemanagementstrategies croppingimpact ofvarious practices of pesticideresistance, andthe host-parasite interactions, evolution models forcomparative studiesof makethemexcellentproperties in thehabitatstheyoccupy. These also canbeenvironmental indicators that govern suchecosystems;they communities subjecttointeractions Especially insoils, theylivein modes according totheirstage). commensal, parasitic, ormixed strategies (e.g.survival free-living, environments andhavevarious species). They are foundinall (26,000 and diverseontheearth group, oneofthemostabundant Nematodes belongtoamonophyletic parasitic nematodes The caseoftheplant- bioagresseurs enmilieutropical, Cirad/ « Productions fruitières» ethorticoles Researchers involved:Researchers PVBMT (Peuplements végétaux et Mireille Fargette(IRD), Mateille(IRD) Thierry UPR Horticulture, Département Researchers involved:Researchers de la diversité descommunautés de ladiversité (Inra, MontpellierSupAgro, Cirad, IRD) Researchers involved:Researchers de nématodesphytoparasites » Philippe Ryckewaert, FrédéricChiroleu, CBGP, Centre deBiologie et Program «Écologie etgestion de GestiondesPopulations Université delaRéunion) Director: Bernard Reynaud, Director: DenisBourguet, [email protected] [email protected] [email protected] Main teams Head: Philippe Vernier, Jean-Philippe Deguine Laurence Arvanitakis UMR 1062 (Flhor, Cirad) UMR C53 Bernard Reynaud, SergeQuilici, Patrice Cadet(IRD), Dominique Bordat, Évolutive enagrosystème maraîcher Contact: Dominique Bordat, of populations pestsandnaturalenemiesnative todifferent geographic regions. includethebiologicalandgenetic differentiationOther topicsunderstudyinthelaboratory select new naturalenemy populationshaving females withgreat reproductive potential. Current research assessesfemale parasitoidbehaviour, dependingonhost patch size, to parasitoids ; andmostrecently vegetables / natural enemiesinseveral systems: vegetables /leafminerparasotoids; cabbages/DBM pollution, grower/consumer networks)? The Laboratory developments, parksmanagement, sharingofresources withthecityandmanagingwater exchange)? How toidentifyandrecognize otherperiurbanagriculturalsystems(land How tofindtoolsfor thefieldofsocialstudies(market access, transfer, technolgy and • To improve thecompetitiveness ofthefieldstudies. • To propose anagroecological production systemandtodefinethequality ofthe The research follows two mainlines: on theenvironment, andimprovement ofmarket access for the growers. residuals invegetable production, reduction ofthenegative impactofvegetable production perishability andseasonality. This fixed topichas three goals: thediminution ofpesticide vegetables, plantainbananas, roots, andtubers, for mostofwhichtheprincipalconcernis environmental protection. products, Regarding horticultural theUPRlimitsitsscopeto production,local self-suffiencyoffood securityandsustainablehorticultural aswell as is toensure sustainableagriculturalproduction intropical andmediterranean areas, including The topicoftheUPR “Horticulture”, from crops» the«FruitandHorticultural Department de nématodesphytoparasites gestion deladiversité descommunautés IRD) withintheprogram « Inra, Montpellier SupAgro, Cirad, the objectivesofCBGP(UMR1062 and environments suchassoils, management ofcomplexhabitats In order toengageresearch on pathogenic effect. without modifyingtheirglobal to specieserosion incommunities against fewspecies. This couldlead to chemicalcontrol, are targeted practices, developedasalternatives rotations, resistance andbiological nematodes are high.Crop very crops duetoplant-parasitic losses ofthetwenty mostimportant economic pointofview, worldyield for nematodecontrol. From an ecological andagronomic challenge nematode communityapproach isan of soilsonplantpathogens, the resistance/resilience capacities nematode interactions andthe Due tothecomplexity ofsoil- functions. as well astounderstandhabitat and theirecologicaladaptation, from tropical areas totheMediterranean Sea Horticultural agroecological management environment? How tomaintainsoilfertility? products. How toreduce nitrateandresidualproducts pesticidesinhorticultural andthe Agroecology: anothervisionof Agroecology: studiestheinter-relations between crop plants, pests, and Écologie et » isto [email protected] B. tabaci of CBGP. the molecularbiologyfacility areExperiments outin alsocarried (15 000slides, 2300types)isstored. collection ofpreserved nematodes in whichtheMNHN/IRDreference autoclave;optics;CIV) waste-water; treatmentand roots; chloride of (nematode extraction from soils managed asatechnicalplatform (government agreement) ofCBGP, the NS2quarantine laboratory Such studiesare outin carried health. utility ofbiologicalindicatorssoil adaptation incommunitiesandthe anthropogenic stresses onspecies more knowledge abouttheeffectof control methods)by producing management (i.e. sustainable of thesestudiesrelates tonematode physiochemical factors). The goal plant interactions, predators, of theirenvironment (e.g. host- biotic andabioticconstraints coexistence, dispersion) andthe history,evolutionary species interspecific relationships (e.g. in communitiesby studying understand assemblageprocesses /parasitoids. Biodiversité entomologique etd’Écologie Biodiversité sustainable agriculture Controlling populations

© M.S. Garcin Agroecological management of pest mites in vineyards

Mites are important This starting-point observation stimulated research carried out pests in vineyards. over more than 10 years in the laboratory of “Écologie Animale et Their impact has Zoologie Agricole” of Agro-Montpellier on the natural colonization greatly increased since of vineyards by Phytoseiidae. These studies showed the 1950 with the massive importance of uncultivated areas bordering plots (for structure, use of pesticides, floral composition), and especially the effect of plant diversity Phytoseiid predatory especially insecticides. on the occurrence, abundance, and diversity of predatory mite with phytophagous Several hypotheses mites. Several plant species were shown to be favourable to mites on a vine leaf have been proposed Phytoseiidae. These studies have also documented wind dispersal to explain these of Phytoseiidae from uncultivated areas to vineyards. However, outbreaks including the factors affecting the settlement of these predators in vineyards development of pesticide have not been well characterized, as several parameters and resistance in pest mite populations and the elimination of natural interactions (e.g. grapevine growth and development, water enemies by broad-spectrum agrichemicals. stress, distance to uncultivated areas, competition between settled and arriving species and populations, leaf structure) In order to decrease pesticide applications, studies on biological may be involved. These studies are still in progress involving control of these mites have been carried out. The most efficient numerous collaborations with professional agricultural institutes natural enemies are predatory mites belonging to the family (Chambres d’Agriculture, ITV). Research concerning the impact Phytoseiidae. These mites are generalist predators feeding on prey of agroecosystem diversity on the structure of the Phytoseiidae but also on other foods, such as pollen when prey are absent. community is also underway. These studies concern the impact Due to this generalist diet, these predators should persist in of hedges and other vegetation bordering fields and also deal fields independent of prey density. However, their development with intra-plot diversity, studies on weed management, and rate (fecundity) is rather low, which prevents their mass-rearing agroforestery (PIRAT Project on the site of Restinclières). The for large-scale releases. Thus, inoculative release strategies were first results from these studies show changes in the diversity of developed as a first step to enhance their occurrence. predatory mites when trees are interspersed with vines.

While success has been observed in some cases, in several others These scientific studies are directly linked to works carried out in these releases did not lead to control of pest mites and the the “Centre de Transfert” of Montpellier SupAgro, concerning side- factors involved are still not known. Studies on other biological effects of pesticides on predators, especially predatory mites, with control strategies have been carried out to enhance the use the common goal of developing an integrated mite management of these efficient predators. Indeed, in some vineyards using program using fewer pesticides and more biological control. integrated pest management (IPM), where pesticides used were selected according to their low side-effects on Phytoseiidae, Contacts: Marie-Stéphane Tixier-Garcin, [email protected] an increase of the numbers of these predators was observed. and Serge Kreiter, [email protected]

© Région Languedoc-Roussillon

VVineyardineyard inin thethe LLanguedoc-Roussillonanguedoc-Roussillon rregion:egion: a highhigh value-value- aaddeddded crop...crop... Biological control Biological 47 48 Biological control Marie-Stéphane Tixier-Garcin (Montpellier SupAgro) (Montpellier Marie-Stéphane Tixier-Garcin Team «Écologie intégrative desSystèmes Team «Écologie animaleetZoologie Olivier Bonato(IRD), Frédéric Pellegrin (IRD), working onthistheme Researchers involved:Researchers Nathalie Gauthier(IRD), Claire Vidal (IRD) (Inra, MontpellierSupAgro, Cirad, IRD) CBGP, Centre de Biologie et Populations-Environnement » Serge Kreiter (Montpellier SupAgro), de GestiondesPopulations Director: DenisBourguet, [email protected] Researchers-teachers involved:Researchers-teachers agricole - Acarologie » Other teams Necrosis of UMR 1062 Jacques Fargues(Inra), similis Radopholus banana tree onroots of © T.Mateille the caseofplant-parasitic nematodes Parasite biodiversityandsoilhealth: root-knot of theeffect ofcropping strategiesnotonly onthemost dangerous species(suchas enhances active biologicalcompetition. Consequently, organicagriculture allows analysis amendments. All thesemethodscontribute toanincrease insoilbiodiversity, which crop diversity, rotations withunsuitablehost plants, green manures, andnatural, organic • cropping systems: Three research paradigmsare defined here: can laterbeconfirmedthrough functional approaches. Statistical methodsusedfor environmental dataanalysis are abletotesthypotheses that understanding ofcommunity structure incropping systemsorfragileenvironments. of environments withcontrastingcharacteristics(i.e. different inputlevels) allows “recovery” capacity ofasoil(soilresistance orresilience). The comparative investigation capacity ofanenvironment tofacilitateepidemicphenomena; iii)the “buffering” and/or on: i)thedisturbancelevel ofanenvironment (e.g. duetohumanactivity), ii)the Diversity andpopulationlevels inanematodecommunity canprovide information approacheshost-parasite binary are notsustainable. susceptibility tomicrobial antagonists (“biocontrol” issue)hasshown thattherapeutic due toparasiticnematodes(“resistance andcrop rotation” issues)andonnematode without any specificdamagebeingallocatedtothem. Research onplantpathology (e.g. In naturalecosystems, speciesdiversity high ofnematodecommunities isvery number ofnematodesinthecommunity. subsequent crops decreases whendiversity increases, withoutany decrease ofthetotal with thelengthoffallow andthattheglobalpathogenesisofcommunities on developed onfallowing showed thatdiversity ofnematodecommunities increases environment (e.g. organicagriculture), nematodediversity isusually higher. Studies plant associationsorcrop rotations, etc.), orincropping systemswhichrespect the dead ends. Inlow-input cropping systems(withextensive agriculture, fallow practices, control techniquesare basedonhighinputsandcanleadtoeconomicecological of competitionandcancausesevere damagetoplants. Inthese cropping systems, frequent abundant. andvery These colonizingspeciesare aggressive very intheabsence than 10plant-parasiticspeciesinthesamesoilsample). Someofthespeciescanbevery Intensive cropping systemsusually produce low-diversity nematodecommunities (less the soilcommunity whoseresistance/resilience capacity shouldbepreserved. here inwhichnematodeswould notbeconsidered only asaggressorsin butaspartners the disequilibriuminducedby high-inputcropping systems. A new paradigmissuggested true for plant-parasiticnematodes. Inthatcontext, pest-control techniquesincrease within-field speciescompositionandfunctionsinpathogencommunities. This isalso High-input agriculturalpracticescanexcaerbateepidemicsituations. They change ≥ 20 speciesinnematodecommunities in Atlantic andMediterranean coastaldunes) Agroecology: anothervisionof Agroecology: Meloidogyne in organicagriculture, nematode-control techniquesinvolve spp.) butonthewhole nematode community, sustainable agriculture ••• and predators). regulations(interspecific competitions) andvertical (from plant this project isto combineresearch onhorizontalregulations directly for food withendoparasiticnematodes. Innovation in On theotherhand, ectoparasiticnematodeswould compete Pochonia chlamydosporia the nematode)by microbial antagonists (e.g. Pratylenchus marham grass( diversity ofregulations. For example, intheduneswhere • low-input andnaturalecosystems: Sahelian vegetable-producing areas” (MAE-DURASgrant). for plant-parasiticnematodemanagementinMediterranean and is beingdeveloped intheproject Nematus “Integrative approach (e.g. approximately 8000hainMorocco). This typeofresearch In Maghreb countries, organicagriculture surfacesexpandfaster areas occupiesabout40%ofthe2000hacultivatedinFrance. in bothProvence-Alpes-Côte d’AzurandLanguedoc-Roussillon examined through acomparative approach. Organicagriculture situations inbothsidesoftheMediterranean Seathatcanbe an increased activityinorganicagriculture withbalanced such research becausetheMediterranean region hasexperienced Mediterranean vegetable cropping systemsare appropriate for ••• nematodes ( the caseofplant-parasitic nematodes Parasite biodiversityandsoilhealth: without biasintroduced by chemicaltreatments. spp.) are controlled by thehost-plant(resistance to Meloidogyne maritima, Heterodera arenaria Ammophila arenaria ) ; andby root symbionts(e.g. mycorhiza). ) wasintroduced, endoparasitic studies reveal agreat Pasteuria penetrans, and Mireille Fargette, [email protected] otcs her Mtil, [email protected], Mateille,Contacts: Thierry for protection/re-establishment of degradedenvironments. strategies for sustainablemanagement ofcropping systemsand on thesoilresilience concept, thisapproach shouldleadto approach provides information onpopulationassemblages. Based (interspecific competitions, biologicalandedaphic constraints) This descriptive (biology, taxonomy, phylogeny) andfunctional and managementofnematodediversity insoilcommunities. This eco-epidemiologicalapproach isfocused onthestudy fonctions danslesécosystèmes des nématodes: populationsàleurs dufonctionnement deleurs developed withintheproject NemaGEco « of nematodecommunities. Research onthissubjectisbeing essential tounderstandingspatialandtemporaldevelopment Knowledge ofpriorusetheland(previous crops, is history) introduction inthe African savannas. example, crop wedges intheLandesforest ornew crop communities are studiedinchanging land-usesituations. For • shiftingsituations: and Serge Morand, [email protected] Patrice Cadet, [email protected] ecologicalfactorsthatstructure nematode Infesting »(CNRS/ADEME-ECOGERgrant). Hoplolaimus pararobustus on aroot ofbanana tree Écologie intégrative , © T.Mateille ,

49 Biological control 50 Biological control in cottonfarming systems populations ofphloem-feedinginsects Agroecological managementof to betaken intoaccountinthe longtermtodevelop durable paradoxically makes itpossible toinventory theprincipalparameters This diagnosis, whichisdifficultbecauseofthenumber ofvariables, 6) by the massive useofinsecticidalproducts inmarket gardens. development attheexpenseof pest-resistance traits; 5) by the moderncultivatedvarieties, withstrong vegetative 4) by the active compoundsusedsuchasthepyrethroids; 3) by thetechniquesoflow-volume pesticidetreatment; fertilizers); (e.g. increased amountsofmineralfertilizer, nitrogenous inparticular methods having ruptured apre-existing balanceintheentomofauna technology, cultivationmethods, and phytosanitary protection increase incottonandmarket-gardening surfaces; evolution of 2) by thechangesinprevalence crops, ofcertain an inparticular the 70’s; inmany therainfalldeficitsobserved tropicalparticular areas since 1) by theevolution oftheabioticfactorsenvironment, in How toexplainthechangeinstatusoftheseinsects? morphological diversity, andhighgeneticvariability. rapid development ofresistance toinsecticides, remarkable share commoncharacteristics, includinghighrateofreproduction, one hosttoanother, according totheconditions. These insectsalso enables themtopassrathereasily from onefieldtoanotherorfrom producing areas oftheworld; they are polyphagous, very which Aphis gossypii natural enemies(predators, parasitoids, entomopathogenicfungi). between theseinsects, theirplantenvironment, andtheirdiverse is theconsequenceofarupture ofthebalanceinitially established advanced thatthechangeinstatusofthesephloem-feeding insects is stimulating research aimedattheircontrol. hasbeen A theory of theworld. Their proliferation isnow regularly and observed Bemisia tabaci two Homopterans, theaphid In cottonculture 1 and (Gennadius), have becomemajorpestsinvarious areas B. tabaci (Gossypium hirsutum) are distributedinmostofthecotton- Aphis gossypii inthelasttwo decades, Glover andthewhitefly Agroecology: anothervisionof Agroecology: biodiversity incottonfarmingsystems. to theconceptofsustainabledevelopment, by enhancing inparticular soil, and thesafeguarding ofindigenous biologicalsystems, according the cropping system, oftheagronomic potentialofthe conservation reference for ensuringtheprofitability andthusthesustainabilityof human healthandbiologicaldiversity). These are true terms of methods thatare theleastdetrimentaltoenvironment (including is ofacurative nature, aimingatimplementation ofpestcontrol within thesamefarmingsystem(conceptofterritory). The lastphase its environment, oreven ofthewholeconcernedplantation(s) The secondphaseaimsatthediagnosisofstateplotand theactionofnaturalenemies.risks asmuch aspossibleandsupport aimstoreducephase ofthisintegratedstrategy thephytosanitary solely through strictrespect toanintegratedprotocol. The initial It isclearthatthesuccessofsuchanapproach cannotbeassured be avoided. level, ofvarioustechniques, amongwhichsyntheticpesticidesshould intervention; theotherrelies oncurative employment, ontheplot the agroecosystem atthelevel ofpreventive organizationand approach totheregulation ofpestpopulationsandthusconsidering different strategies: one, regarded asapriority, favoring an ecological agriculture, whichendeavors toreconcile theadvantagesoftwo concept ofintegratedprotection inthecontextofsustainable for thisagroecological process isproposed. Itisbasedonthe From theobservations, experiments, andreflection, thefoudation of balance. adopt new procedures andpractices, allowing areturn toasituation to avoid anaggravation ofthesituationand, ontheotherhand, to is, ontheonehand, tobreak withpastpracticesorusagesinorder strategies for managementofinsectpestpopulations. The objective [email protected] Contact: Jean-Philippe Deguine, sustainable agriculture 2 © J.P. Deguine © J.P. Deguine Aphis gossypii 2. African cottonfield 1. 3. and adultsofcottonaphids a ladybird predator oflarvae 3 Winged adultsof of Larva Harvesting inan © J.P. Deguine Cheilomenes vicina, 51 Biological control © Cirad Dissemination of scientific and technical culture

The Common Resource Documentation Center (CCRD) houses the resources (print and electronic resources) of three units of C.I.L.B.A (Complexe International de Lutte Biologique Agropolis) related to biological control and management of pest populations. Biological control Biological

52 documents (booksandjournals) enabling interrogation ofauniqueinterfaceandlocationthe than 50librariesandunitsoftheLanguedoc-RoussillonRegion, Roussillon public Internet, the«B FSTA, Medline, Biosis, andZoologicalRecord. Also available on Abstracts, Agricola, CCConnect, Econlit, Web ofScience, databases are now available inthethree unitsincluding: CAB • Provide accesstoexternal databases: developments, andnew electronic resources • Circulate information onacquisitions, websites ofCBGPandCILBA publications ofthescientists)andmake themavailable onthe • Develop commoncatalogues searches, loans, interlibrary interrogation ofdatabases, etc. Contents, documentdelivery, training, retrospective computer labs: lendingservices, reference services, profiles onCurrent • Provide arange ofservices • Coordinate ∆ ∆ ∆ ∆ ∆ ∆ ∆ and internationalcollaborations • Develop anetwork ofnational ∆ T

and National (Beltsville,Agricultural Library Maryland, USA) (CSIRO,Mountain Library Canberra) institutions Montpellier SupAgro) inMontpellier(Inra, ofallthepartners services IRD, Cirad, collaborations et tropicale international (IAALD). Member ofthe “IFR Network ofINRAlibrarians European (EBCL, BiologicalControl Laboratory Montpellier) CSIRO (Montpellier)andtheBlack European Laboratory Network ofthelibrariansall Agropolis International Documentation centers, communication andedition Developing anetwork ofnationalandinternational Professional associations, bothFrench (ADBS)and aims to: Center he (BOMLR) », aglobalfree-access catalogueofmore Common Resource Documentation ,” Montpellier themanagementofresources (CCRD), headedby ChristineSilvy, ibliothèque Ouverte MontpellierLanguedoc- ibliothèque Ouverte Biodiversité continentale,Biodiversité méditerranéenne totheusersofthree (http://www.bomlr.info (e.g. journals, books, [email protected] most ofthese Christine Silvy, technological ). 53 Biological control 54 Biological control Denis Bourguet Thematic «Systématique-Taxonomie, Phylogénie-Phylogéographie » (Inra, Montpellier SupAgro, Cirad, IRD) UMR 1062CBGP, Centre deBiologie etdeGestiondesPopulations Denis Bourguet Program «Écologieetgestiondeladiversité descommunautésdenématodesphytoparasites » (Inra, Montpellier SupAgro, Cirad, IRD) UMR 1062CBGP, Centre deBiologie etdeGestiondesPopulations Denis Bourguet Group «Processus évolutifs etgestiondesbio-agresseurs » Team «Écologieintégrative desSystèmes Populations-Environnement » (Inra, Montpellier SupAgro, Cirad, IRD) UMR 1062CBGP, Centre deBiologie etdeGestiondesPopulations Denis Bourguet Team «Écologieintégrative desSystèmes Populations-Environnement » (Inra, Montpellier SupAgro, Cirad, IRD) UMR 1062CBGP, Centre deBiologie etdeGestiondesPopulations Miguel Lopez-Ferber (EMA) LGEI Laboratoire Géniedel’Environnement etnaturels industriels etdesrisques industriel Denis Bourguet Team «ÉcologieanimaleetZoologie agricole-Acarologie » (Inra, Montpellier SupAgro, Cirad, IRD) UMR 1062CBGP, Centre deBiologie etdeGestiondesPopulations Andy Sheppard/Mic Julien Thematic «Bioagresseurs émergentsetbio-invasions » CSIRO, Laboratoire européen Pascal Marnotte (Cirad) Unité deRecherche «Systèmes canniers» Walker Jones Team «Lutte biologiquecontre lesbio-invasions » USDA-ARS, EBCL, Laboratoire Européen deLutte Biologique Walker Jones Team «Quarantaine » USDA-ARS, EBCL, Laboratoire Européen deLutte Biologique Walker Jones Team «Unité d’évaluation génétique» USDA-ARS, EBCL, Laboratoire Européen deLutte Biologique Jean-Dominique Lebreton (UMR 5175,CNRS,UM1,UM2,UM3, Montpellier SupAgro, Cirad) CEFE, Centre d’écologie fonctionnelleetévolutive Jean-Christophe Auffray Team «GénétiqueetEnvironnement » (CNRS,UM2) UMR Institut desSciences del’Évolution Philippe Vernier «BiodiversitéLaboratory entomologique etd’Écologie Évolutive enAgrosystèmes maraîchers » (Flhor, Cirad) UPR Horticulture, «Productions Département fruitières » ethorticoles Bernard Reynaud (Cirad/Université delaRéunion) UMR C53PVBMT (Peuplements végétauxetbioagresseurs enmilieutropical), appearance inthisdocument. R following inorder chart of are mentionedonthe esearch unitsandteams em 1 Teams Topics covered by c. Legislationandregulation aspects: b. Considering invasive species a. The taxonomic tool in agro-ecosystems 1. quarantine issues organisms,harmful agreements and helping biologicalcontrol Characterizing biodiversityCharacterizing research teams •• •• •• •• •••• •• •••••• • •• •• a c. Agroecology, anothervisionofsustainable b. Sustainable plantprotection a. Classicalbiologicalcontrol: screening and 3. Controlling pestpopulations b. Characterization ofgeneticdiversity a. Evaluation ofhostspecificityandmulti- biodiversity anditsinteractions 2. Understanding andevaluating agriculture for sustainableagriculture introducing natural enemies trophic interactions 1 ••• •••• ••••• • •• •• ••• b 1 • •••• c 2 ••• a 2 b (July 2006) (July 3 ••• a 3 b 3 c in thedomainofBiological control at Agropolis International Education andtraining The training-education programme as vocationaltraining institutions). and schoolsofengineers(aswell member institutions, universities programme delivered through its A Montpellier SupAgro, Montpellier SupAgro Bac +5 Bac +8 Bac +3 ee degrees title level institutions training-education proposes acomplete gropolis International Inra (BSc withprofessional professionnelle professional scope) professionnel “ recherche Ingénieur Doctorat (MSC with Licence Master Master degree scope) (PhD) (MSc) ” speciality Ecology, Bio-diversity andEvolution Evolution and Agronomical important insects:practiceAgronomical important inidentifi Acarids, pestsandhelpersforplants:presentation andidentifi Integrated systemsbiology, agronomy andenvironment Biology, Geo-science, Agro-resources andEnvironment speciality 'Agronomy andIntegrated systems' farming Phyto-genetic resources andbiologicalinteractions courses related tothisdomain. below detailthetraining-education elaborated onrequest). The tables training modules(existingor master, PhD), aswell asvocational degree,technician, engineering courses” (from Bac +2toBac +8: morecomprises than 80 “diploma Crop Protection andEnvironment Sustainable crop production Agronomy andagro-food Integrated farming Agro-horticulture Ecology pathway Vine andwine Vine Plant health title Courses and cation in laboratory (5days) cation inlaboratory the training isdelivered. trainings andtheinstitutionswhere diplomas, ofthe adescription They specifythelevelsof Short cation (4days) Agrocampus-Rennes, Montpellier SupAgro, Montpellier SupAgro, Montpellier SupAgro, Montpellier SupAgro, Montpellier SupAgro, Montpellier SupAgro INRA Paris-Grignon UM1, UM2,UPVD Castelnaudary AgroParisTech UPVD, LEGTA degrees courses INH Angers institutions UM2 55 Biological control Glossary

Agroecology: study of correlations Co-evolution: the evolution of two Haplotype: a contraction of the between plants, , man and or more interdependent species, each phrase “haploid genotype”: the genetic environment within agricultural adapting to changes in the other. It occurs, constitution of an individual chromosome. ecosystems. for example, between predators and prey and between insects and the flowers that Inoculative (classical) biological Allochtonous species: species which they feed on or pollinate. control: importation and release of are outside of their natural region biological control agents into an area usually as result of an intentional or Complex of species: a group of closely in which they are not already present, accidental dispersion by human activities related taxa whose taxonomic or genetic with intent to establish permanent, (also known as an introduced or exotic differentiation is difficult to resolve with self-sustaining, and self-disseminating species). confidence based on current data. populations of the agent.

Allopolyploidy: polyploidy resulting Cryptic species: distinct species that are Intraspecific variability: variability from an interspecific cross (thus extremely similar morphologically. arising or occurring within a species. containing two different genomes). Diagnosis: determination of the nature of a Inundative biological control: release Barcoding: method of taxonomic pathological condition ; recognition of a disease. of large numbers of a biological control identification of eukaryotic organisms agent relative to the number of the target based on the diagnostic power of small Diploid: a full set of genetic material, species, in expectation of a rapid effect. fragments of DNA. consisting of paired chromosomes; one There is no implication that the released chromosome from each parental set. biological control agent is of exotic origin Bioindicators: species which, due or will establish a permanent population. to their sensitivity to environmental Entomopathogenic: acting as a parasite of conditions, provide a precocious insects and killing or causing serious disease. Invasive species: any species, including indication of biotic or abiotic its seeds, eggs, spores, or other biological perturbations of the environment, Epizootic: a disease epidemic in an material capable of propagation, that is often due to human activities. animal population. not native to an ecosystem and whose introduction does or is likely to result Biopesticides: pesticides originating from Flow cytometry: a technique making it in rapid, uncontrollable spread and natural substances or from living organisms. possible to sort particles, molecules, or cells at population growth of that species. high speed using a laser beam. The re-emitted Biotype: a group of organisms having the light (by diffusion or fluorescence) makes it ITS (Internal Transcribed Spacers): same genotype. possible to classify a group according to several RNA fragments used as molecular criteria and to sort and count each subgroup. markers. Cluster: a group of the same or similar elements gathered or occurring closely Genetic distance: a way of measuring Karyology: the study of chromosome, together. the amount of evolutionary relatedness physiology, including their shape, relative between two species or distinct size, banding patterns, number, etc. Co-dominant: describing a genetic populations of a species by counting the marker for which it is possible, for a given number of allelic substitutions or point Microsatellites, or Simple Sequence locus, to distinguish either homozygous mutatations that have arisen in each Repeats (SSRs): polymorphic loci Biological control Biological condition as well as the heterozygous population since their divergence from a present in nuclear DNA that consist of 56 condition. common ancestor. repeating units of 1-4 base pairs in length. They are typically neutral, co-dominant Phylogeography: the study of the RFLP (restriction fragment length (q.c.) and are used as molecular markers geographic distribution of genetic polymorphism): genetic variation which have wide-ranging applications in lineages through construction of between individuals measured by the field of genetics, including kinship, genealogies of populations by comparing comparing differences in DNA fragment selective breeding, and population studies. their DNA sequences. size resulting from sequence differences at restriction enzyme recognition sites. Morphometry: measurement of the Polyploidy: the condition of some form of an organism. organisms of containing more than one Sequencing: determination of the order set of paired homologous chromosomes of nucleotides (base pairs) in a DNA or PCR (polymerase chain reaction): per cell. RNA molecule or the order of amino acids a molecular biology technique, for in a protein. enzymatically replicating DNA in vitro. Resilience (in ecosystems): The technique allows a small amount a measure of how much disturbance Taxon: a taxon (plural taxa), or taxonomic of a DNA molecule to be amplified (e.g. from storms, fire and pollutants) unit, is a grouping of organisms (named exponentially. Since it is an in vitro an ecosystem can handle before shifting or unnamed) based on relatedness. Once technique, it can be performed without to a qualitatively different state. It is the named, a taxon will usually have a rank restrictions on the form of DNA (apart capacity of a system to withstand shocks (e.g. species, genus, family, subfamily) and from size limitations) and it can be and surprises and then rebuild itself. It is can be arranged within a hierarchy of extensively modified to perform a wide during this rebuilding phase that renewal related organisms. array of genetic manipulations. and innovation take place in resilient systems. Taxonomy: was once only the science Phylogeny: the study of the geographic of classifying living organisms, but later the distribution of genetic lineages word was applied in a wider sense, and through construction of genealogies of may also refer to either a classification of populations by comparing their DNA things, or the principles underlying such sequences. classification.

© F. Hérard Dolichomitus messor (Hym. : Ichneumonidae), a parasitoid known from Lamia textor (Coleoptera : Cerambycidae) in Europe. Biological control Biological 57 58 Biological control DURAS project DRAF/SRPV USDA/ARS C.I.L.B.A. FDGEON ECOGER ENGREF CAREPS URCAM BOMLR ADEME PVBMT DAISIE INA PG MNHN ESALQ TYLCV LEGTA CSIRO IAALD CIRES MSIRI PIRAT CITES CTIFL UPVD UMIII SASRI CBGP DSCP CNRS ADBS LNPV Cirad EBCL CEFE SRPV CETA ACTA DESS IRAD UMII BSES UMR DBM CIPV LGEI Isem IRTA MAE EMA UPR UMI DAA DEA Inra LOA IPM CEE INH FAO ARS IRD COI PBI IFR ITV DU Bureau ofSugar Stations Experiment Bibliothèque Ouverte Montpellier Languedoc-Roussillon Agricultural Research Service Agence del’Environnement etdelaMaîtrise del’Énergie desprofessionnels del’informationetladocumentation Association decoordinationAssociation techniqueagricole Convention surlecommerce internationaldesespècesdefauneetfl Centre Inter-organisme deRecherche etd’Expertise enSystématique pour ledéveloppement Centre decoopération internationaleenrecherche agronomique Convention Internationale pourlaProtection des Végétaux Complexe International deLutte Biologique Agropolis Collège d’Enseignement Technique Agricole Centre d’Écologie Fonctionnelle etEvolutive Communauté ÉconomiqueEuropéenne Centre deBiologie desPopulations etdeGestion Centre Rhône-Alpes d’Epidémiologie etdePrévention Sanitaire Centre Technique Interprofessionnel desFruits etLégumes Cytochrome OxidaseSubunit I Centre nationaldelarecherche scientifi Ministère desAffaires étrangères Lettre Offi Laboratoire National delaProtection des Végétaux et desrisquesindustrielsnaturels Laboratoire Géniedel’environnement industriel Lycée d’Enseignement Général etTechnologique Agricole Institut dessciencesdel’évolution deMontpellier Centre Technique Interprofessionnel dela Vigne etdu Vin El Instituto deInvestigación yTecnología Agroalimentarias Institut deRecherche pourleDéveloppement Institut deRecherche AgricolepourleDéveloppement Insect Pest Management Institut National d’Horticulture Institut nationaldelarecherche agronomique Institut nationalagronomique Paris-Grignon Institut Fédératif deRecherche International Association ofAgricultural Information contre lesOrganismes Nuisibles Fédération desGroupements Départementale deDéfense Food andAgricultural Organization Escola Superior deAgricultura Luiz deQueiroz École Nationale duGénieRural desEauxetForêts (AgroParisTech) École Nationale Supérieure desTechniques Industrielles etdesMines d’Alès European Biological Control Laboratory Écologie pourlagestiondesEcosytèmesetdeleursressources UniversitaireDiplôme Promoting Sustainable Development inAgricultural Research Systems Double Strand Conformation Polymorphism Service RégionaldelaProtection des Végétaux Direction Régionaledel’Agriculture etdelaForêt, d’ÉtudesDiplôme Supérieures Spécialisées d’ÉtudesDiplôme Approfondies AlienInvasiveDelivering Inventories forEurope Diamond Black Moth d’AgronomieDiplôme Approfondie Commonwealth Scientifi United States of Agriculture/Agricultural Department Research Service Union Régionale desCaisses d’Assurance Maladie Unité Propre de Recherche Université dePerpignan Via Domitia Unité Mixte deRecherche Université Montpellier 3 Université Montpellier 2 Université Montpellier 1 Tomato YellowCurl Virus Leaf Service RégionaldelaProtection des Végétaux South Sugarcane African Research Institute Peuplements végétauxetbioagresseurs enmilieutropical Programme Intégré deRecherches enAgroforesterie àRestinclières Protection Biologique Intégrée Mauritius Sugar Research Industry Institute Museum nationald’Histoire naturelle cielle d’Autorisation List ofabbreviations c andResearch Organisation (France) que ore sauvages This publication receives support from the State and the Languedoc-Roussillon Region Research units, member organisations and partners of Agropolis International involved in the field of Biological control...

CBGP www.montpellier.inra.fr/CBGP CEFE www.cefe.cnrs.fr Cirad www.cirad.fr Csiro www.csiro.au EMA www.ema.fr Inra www.inra.fr IRD www.ird.fr Isem www.isem.cnrs.fr Montpellier SupAgro www.supagro.fr UMI www.univ-montp1.fr UMII www.univ-montp2.fr UMIII www.univ-montp3.fr UPVD www.univ-perp.fr USDA www.ars-ebcl.org

In the same collection September 2007 • 60 pages French and english version

Director in chief: Henri Carsalade

Scientific coordinator: René Sforza

English version editor: Brian Rector (EBCL)

Science writer : Isabelle Amsallem (Agropolis Productions)

Participed to this issue: Cindy Adolphe, Laurence Arvanitakis, Marie-Claude Bon, Olivier Bonato, Dominique Bordat, Denis Bourget, Patrice Cadet, Dominique Coutinot, Jean-Philippe Deguine, Gérard Delvare, Michel Ducamp, Pierre Ehret, Mireille Fargette, Jacques Fargues, Régis Goebel, Franck Hérard, Serge Kreiter, Christian Langlais, Miguel Lopez-Ferber, Arnaud Martin, Jean-François Martin, November 2005 • 56 pages Thierry Mateille, Sandrine Maurice, Alain Migeon, Serge Morand, Maria Navajas, Isabelle Olivieri, French and english versions French and english versions French and english versions Jean-Yves Rasplus, Yves Savidan, René Sforza, Andy Sheppard, Christine Silvy, Marie-Stéphane Tixier-Garcin, Claire Vidal Les dossiers d’Agropolis International Illustrations: We like to thank “Les dossiers d’Agropolis International” series is one of the productions of Agropolis the authors and Cirad, Den Colong, Mireille de Freitas, International within the framework of its mission for the promotion of the scientific Gilberto de Moraes, Alain Dutartre, Denise Navia, community competences. Mark Volkovitch, Région Languedoc-Roussillon Each case is dedicated to a broad scientific theme. They comprise a synthetic presentation, Corrections: Isabelle Amsallem, Brigitte Cabantous, easily accessible for all the laboratories and research teams on site working on the theme Marie-Claude Kohler, René Sforza, Christine Silvy, concerned. Nathalie Villeméjeanne The objective of this series is to enable our different partners to gain a better understanding Layout and computer graphics: and a better knowledge of the competence and of the potential available in this community, Olivier Piau, [email protected] but also to facilitate the contacts for the development of scientific and technical exchanges Print: Les Petites Affiches (Montpellier, France) and cooperations. Photo cover: © R. Sforza ISSN: 1628-4240 • Copyright: September 2007