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Forest entomology To: A. H. MacAndrews Who introduced me to the world of forest insects Forest entomology A global perspective By William M. Ciesla Thiseditionfirstpublished2011(cid:2)2011byWilliamM.Ciesla BlackwellPublishingwasacquiredbyJohnWiley&SonsinFebruary2007.Blackwell’spublishingprogramhasbeen mergedwithWiley’sglobalScientific,TechnicalandMedicalbusinesstoformWiley-Blackwell. Registeredoffice: JohnWiley&SonsLtd,TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK Editorialoffices: 9600GarsingtonRoad,Oxford,OX42DQ,UK TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK 111RiverStreet,Hoboken,NJ07030-5774,USA Fordetailsofourglobaleditorialoffices,forcustomerservicesandforinformationabouthowtoapplyforpermissionto reusethecopyrightmaterialinthisbookpleaseseeourwebsiteatwww.wiley.com/wiley-blackwell. TherightoftheauthortobeidentifiedastheauthorofthisworkhasbeenassertedinaccordancewiththeUKCopyright, DesignsandPatentsAct1988. Allrightsreserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystem,ortransmitted,inanyform orbyanymeans,electronic,mechanical,photocopying,recordingorotherwise,exceptaspermittedbytheUKCopyright, DesignsandPatentsAct1988,withoutthepriorpermissionofthepublisher. Designationsusedbycompaniestodistinguishtheirproductsareoftenclaimedastrademarks.Allbrandnamesandproduct namesusedinthisbookaretradenames,servicemarks,trademarksorregisteredtrademarksoftheirrespectiveowners. Thepublisherisnotassociatedwithanyproductorvendormentionedinthisbook.Thispublicationisdesignedtoprovide accurateandauthoritativeinformationinregardtothesubjectmattercovered.Itissoldontheunderstandingthatthepublisher isnotengagedinrenderingprofessionalservices.Ifprofessionaladviceorotherexpertassistanceisrequired,theservices ofacompetentprofessionalshouldbesought. LibraryofCongressCataloging-in-PublicationData Ciesla,WilliamM. Forestentomology:aglobalperspective/byWilliamM.Ciesla. p.cm. Includesbibliographicalreferencesandindex. ISBN978-1-4443-3314-5(cloth) 1. Forestinsects. I. Title. SB761.C5372011 634.9'67—dc22 2010048037 AcataloguerecordforthisbookisavailablefromtheBritishLibrary. Thisbookispublishedinthefollowingelectronicformats:ePDF9781444397871;WileyOnlineLibrary9781444397895; ePub9781444397888 Setin9/11PhotinaMTbyThomsonDigital,Noida,India 1 2011 Contents Acknowledgements,vii 9. BARKANDAMBROSIABEETLES,173 Theauthor,ix 10. LARGECAMBIUMANDWOODBORING INSECTS,203 Preface,xi 11. SUCKINGINSECTS,233 1. THEWORLD’SFORESTSANDTHEIR DYNAMICS,1 12. GALLINSECTS,253 2. FORESTINSECTDYNAMICS,15 13. TIP,SHOOTANDREGENERATION INSECTS,273 3. FORESTINSECTANDHUMAN INTERACTIONS,31 14. INSECTSOFTREEREPRODUCTIVE STRUCTURES,295 4. MONITORINGFORESTINSECTS,THEIR DAMAGEANDDAMAGEPOTENTIAL,49 15. INSECTSOFWOODINUSE,313 5. FORESTINSECTMANAGEMENT,73 References,329 6. FORESTINSECTORDERSAND SubjectandTaxonomicIndex,355 FAMILIES,97 Hostindex,371 7. FOLIAGEFEEDINGINSECTS– Colour plate sections appear between pp. 100–101, LEPIDOPTERA,113 132–133,180–181,228–229and276–277 8. OTHERFOLIAGEFEEDINGINSECTS,149 COMPANIONWEBSITE Thisbookhasacompanionwebsite: www.wiley.com/go/ciesla/forest withFiguresandTablesfromthebookfordownloading Acknowledgements A half-century of work as a forester, forest entomolo- RainierPo€hlmann,DirectorofPublicAffairs,Bavari- gist, remote sensing specialist and forest health pro- anNationalPark,Germany,sharedhisinsightsonbark grammanagerisreflectedinthepagesofthistext.My beetles,forestdynamicsandpeopleandhowtheyaffect chosenprofessionhasprovidedmewithopportunities managementpolicyofanationalpark. to work and travel across much of the USA and also Many friends and colleagues provided photos of Mexico,portionsofCentralandSouthAmerica,Europe, subjects not available from my own archives. These Asia and Africa. I have also had the good fortune to included Ronald F. Billings, Texas Forest Service, meet many outstanding people along the way who S. Sky Stephens, Colorado State Forest Service, became my mentors, colleagues and friends. Aubrey E. Richard Hoebecke, Cornell University, Paula H. MacAndrews, Professor of Forest Entomology at Klasmer, Instituto Nacional de Tecnologia Agrope- the State University of New York, College of Environ- cuaria,Argentina,DavidCappaert,Michigan,R.Scott mental Science and Forestry at Syracuse University Cameron, International Paper Company, Larry introduced me to the world of forest insects when I Barber, Sheryl Costello, Jerald E. Dewey, Jose was an undergraduate forestry student. As a young Negrón and Brian Howell, USDA Forest Service. forestentomologist,Ihadthegoodfortunetoworkwith Severaloftheimagesinthistextwereaccessedviathe people like Gene Amman, A.T. Drooz and Hoover www.forestryimages.org website of the University of Lambert, USDA Forest Service. Robert C. Heller, Georgia,whichistheworkofExtensionEntomologist, USDAForestServiceandlater,UniversityofIdaho,first G. Keith Douce. William Jacobi, Department of introduced meto coloraerialphotos asa toolto map BioagriculturalSciencesandPestManagement,Color- and assess forest insect damage. Russell K. Smith, adoStateUniversity,madeavailablesamplesofdamage USDA Forest Service, provided me the opportunity, caused by ambrosia beetles, carpenter ants, termites guidanceandconfidencetomanageforesthealthpro- and wood borers, photos of which are included in grams in several Forest Service regions and at the thistext. national level. In partnership with contemporaries, The team at Wiley-Blackwell took the title of this including J. W. Barry, W. H. Klein, W. B. White, work “A Global Perspective”seriously. They arranged F.P.Weber,J.D.Wardandmanyothers,weevalu- for constructive and supportive peer review of the atedandintegratedremotesensing,geographicinfor- manuscript by Robert G. Foottit, Agriculture and mation systems and the technology of pesticide Agri-Food Canada, Ottawa, Ontario, Canada. Ward applicationintoforesthealthprotectionprograms. Cooper, Senior Commissioning Editor and Delia Jorge Macias (Mexico), Attilio Disperati, Edson Sandford, Managing Editor of Wiley-Blackwell, Tadeu Iede, Yeda Oliviera and Augusta Rosot Oxford,UKprovidedoverallleadershiptotheproduc- (Brazil), Aida Baldini Urrutia, Osvaldo Ramierez tion and Kelvin Matthews, Production Manager, andPatricoOjeda(Chile),JosephMwangi(Kenya), also in Oxford, arranged for peer review and design Muhammad Yousuf Khan and Hafeez Buzdar ofthecover.MaggieBeveridge,currentlyresidingin (Pakistan), Zhou Jian Sheng and Wang Gaoping Lusaka,Zambia,servedascopyeditorandwasinstru- (China), Cernal Akesen and Musa Erkanat mental in transforming astill unfinished draft into a (Cyprus), Heinrich Schmutzenhofer and Edwin polished document, ready for printing. Jessminder Donaubauer(Austria),MihaiBarca(Romania)and Kaur, Production Editor, Wiley-Blackwell, Singapore, manyotherinternationalcolleaguesintroducedmeto supervisedthelayoutandprinting.PrakashNaorem, the cultures and forests of their respective countries. ThomsonDigital,Noida,Indiaprovidedleadershipto viii Acknowledgements thetypesettingandlayout.Therefore,thisprojectwas Cyprus, Kenya, Indonesia, the USA and many other indeedoneof“global”proportions,withpeoplerepre- places and provided encouragement and support as senting six countries and four continents working Iresearchedandwrotethistext.Severalofherphotos togethertomakeitareality. arealsoincluded. Finally,specialthanksgotomywifeandlifepart- ner, Pat Ciesla, who spent many long days with WilliamM.Ciesla me during field assignments in Brazil, China, Chile, FortCollins,Colorado,USA The author William M. Ciesla earned a Bachelor of Science OfficerfortheFoodandAgricultureOrganizationofthe Degree in Forestry from the State University of UnitedNationsinRome,Italyandin1995,chartered NewYork,CollegeofEnvironmentalScienceandFor- ForestHealth ManagementInternational, aninterna- estryatSyracuseUniversityin1960.Helaterspecial- tional forest health consulting service based in Fort ized in forest entomology at the same institution and Collins,Colorado.Hehasworkedwithforestinsectsin earnedaMasterofScienceDegreein1963.Hebegan over 30 countries and is author or co-author of over his professional carrier in Asheville, North Carolina 160publications.HeisalsoauthorofStranieri:AnItalian with USDA Forest Service and subsequently held Odyssey, a light hearted account of life and travels in assignmentsasaforestentomologistandforesthealth Italy. In 2005 he received the Western Forest Insect programmanagerinseverallocationsthroughoutthe Work Conference - Founder’s Award for outstanding USA.From1990to1995heservedasForestProtection contributionstoforestentomologyintheWest. Preface Insectsarebyfarthemostdominantgroupofanimals consideredmajorforestpestsandtheirpreventionand ontheEarth’ssurface.Overamillionspecieshavebeen controlisanintegralpartofforestmanagement.World- describedandnewspeciesarecontinuouslybeingdis- wide,approximately68millionhaofforestssuffersome covered.Theyareanintegralpartofthebiodiversityof type of insect damage each year. This is more than ourplanetandpresentlycompriseabout80–85%ofthe twicetheareaburnedbywildfires(FAO2005). totalnumberofanimalspeciesknowntoscience.Some Thistextexaminesforestinsectsinaglobalcontext expertsestimatethatthetotalnumberultimatelyrec- and addresses the species of major concern in the ognized may eventually approach 30 million species. world’sforestecosystems.Itisdividedintotwoparts: Morethan1000speciescanoccurinarelativelysimple Chapters 1–5 examine the role of insects in forests, ecosystemsuchasabackyardandcannumbermany their dynamics and their effects on natural forests, millionsperhectareoflandsurface. plantations,agroforestrysystems,urbanforests,wood Inadditiontobeingabundant,insectsareextremely and non-wood products. Approaches to forest insect diverse and adaptable organisms. They are found in monitoringarereviewedandalternativesformanage- virtually every ecosystem. Insects are classified into mentofdamagingforestinsectswithintheframework about30orders.Eachorderisfurthersubdividedand of integrated pest management (IPM) are presented. classified into families, genera and species. In North Thebasisforclassificationofforestinsectsintoorders America alone, some 698 families, with an estimated andfamiliesisreviewedinChapter6.Chapters7–15 95,553speciesarerecognized(Erwin1982,Chapman addressthemainthrustofthistextandprovidedescrip- 2006,Triplehorn&Johnson2005). tions of important forest insects, their distribution, Insectsareanintegralpartofalloftheworld’sforest hosts,lifehistoriesandeconomic,socialandecological ecosystems. Many species serve beneficial and even impacts. These chapters are organized according to critical functions in forests. Some visit flowers and the damage the insects cause, rather than by taxo- pollinateplants.Othersfunctionasagentsinthebreak- nomicgroups. down of dead vegetation. Insects, such honeybees or Thistextiswrittenforforesters,forestentomologists the lac insect, provide products beneficial to humans. and forest health specialists engaged in management Others kill old, mature trees and make way for the and protection of forests, worldwide. It is intended to establishment of young, more vigorous forests. When callattentiontotheimportanceofinsectsintheworld’s insectsbecomeexcessivelyabundant,theycandamage forestecosystems,theneedtoconsidermanagementof treesandforestsandthuscompetewithhumansforthe insect pests as an integral part of sustainable forest goodsandservicesthatforestsprovide.Somegroupsof managementandtoserveasaguidetotheidentifica- insects, such as foliage feeders and bark beetles, are tionoftheworld’smajorpestspecies. Chapter 1 The World’s Forests and their Dynamics INTRODUCTION world’sforests presently store an estimated240 giga- tonnes(Gt)ofcarboninwoodybiomassandatotalof Forestscover3.952billionhaor30.3%oftheEarth’s 683Gtofcarboninforestecosystemsasawhole(FAO surface.Otherwoodedlandscoveranother1.3billion 2005,2009a). ha.1 They provide habitat for many living organisms. The world’s forests have been subject to human Moreover,theyprovideawealthofgoodsandservices pressuresincethebeginningofcivilization.Largeareas forhumans.Forestsareasourceofbothwoodandnon- have been deforested to make room for agriculture, woodproducts,includinglumber,pulpwood,fuelwood, communities, industrial sites, roads and highways. resinandfooditemssuchasnuts,fruits,mushrooms, Additional areas have been degraded as trees of the edibleplantsandgame.Inaddition,theyprovidepro- most desirable species and quality were harvested, tective cover for watersheds, range for domestic ani- forestswereovergrazedbydomesticlivestockorburned mals and are an important source of recreation and for landclearingorto drivegame.Presently, the rate spiritual refreshment. A more recently appreciated of forest loss due to land use change is estimated at valueofforestsistheirabilitytoremoveexcesscarbon 13millionha/year,resultinginanetreductionofforest from the Earth’s atmosphere, much of which is pro- area of about 7.3 million ha/year. Only 36% of the duced by humans through burning of fossil fuels or world’sforestsareregardedas“primaryforests.”These clearingofforests,andstoreitinwoodybiomass.The aredefinedasforestsofnativespeciesinwhichthereis no clearly visible indication of human activity and ecological processes are not significantly disturbed 1Forestsaredefinedaslandspanning0.5hawithtreeshigher (FAO2005,2009a). than5mandacanopyofmorethan10%.Otherwoodedland isdefinedaslandspanningmorethan0.5hawithtreeshigher Forestshavethecapacitytoregenerateandproduce than5mandacanopycoverofbetween5%and10%(FAO goodsandservicesforhumansonacontinuingbasis, 2005,2009a). provided they are managed in a sustainable manner. ForestEntomology:AGlobalPerspective,FirstEdition.WilliamM.Ciesla. (cid:2)2011WilliamM.Ciesla.Published2011byBlackwellPublishingLtd. 1

Description:
Content: Chapter 1 The World's Forests and Their Dynamics (pages 1–14): Chapter 2 Forest Insect Dynamics (pages 15–30): Chapter 3 Forest Insect and Human Interactions (pages 31–47): Chapter 4 Monitoring Forest Insects, Their Damage and Damage Potential (pages 49–72): Chapter 5 Forest Insect
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