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Insects as biotic disturbance agents Allan Carroll Associate Professor Lecture outline/goals: o Forest disturbances: abiotic versus biotic o Insects as biotic disturbance agents o Insects and “forest health” in the headlines o Forest health: what it really means o Insects and forest disturbance o Examples: the major groups © A.L. Carroll Wildfire, wind, etc. Abiotic “Healthy” forests require change or disturbance © D. Powell, USDA Forest Service, Bugwood.org Disturbance = any discrete event that disrupts forest ecosystem, community, or population structure and/or function Natural disturbance agents comprise mainly… Insects & disease Biotic © A.L. Carroll © A.L. Carroll Timber volume (m3 × 106) lost per year Relative impacts of abiotic versus insect disturbance in Canadian forests 100 Insects have much greater impacts than wildfire Disturbance annually affects many thousands of hectares 50 High public profile o “Forest health” 0 Source: National Forestry Database (http://nfdp.ccfm.org/) © A.L. Carroll Insects and “forest health” in the news: the emerald ash borer “Scientists Race to Prevent Ash Tree’s Extinction”, New York Times, Oct. 8, 2009 © D. Cappaert, Michigan State Univ., Bugwood.org © J. O’Brien, USDA Forest Service, Bugwood.org © J.W. Smith, USDA APHIS., Bugwood.org © D. Cappaert, Michigan State Univ., Bugwood.org Insects and “forest health” in the news: the Sirex wood wasp “Alien Wasp Kills Whole Forests”, ABC News, Mar. 8, 2006 © W. Cranshaw, Colorado State Univ., Bugwood.org © D. Haugen, Bugwood.org © D. Haugen, Bugwood.org © D. Haugen, Bugwood.org Insects and “forest health” in the news: the spruce beetle “Billions of Beetles, Wildfire Spread Imperil Northern Forests”, Associated Press, Aug. 23, 2009 © E.H. Holsten, USDA Forest Service, Bugwood.org © W.M. Ciesla, Forest Health Mgmt Int, Bugwood.org © A.S. Munson, USDA Forest Service, Bugwood.org © E.H. Holsten, USDA Forest Service, Bugwood.org Insects and “forest health” in the news: the mountain pine beetle © A.L. Carroll “Beetles Take a Devastating Toll on Western Forests”, New York Times, Jul. 13, 2004 © A.L. Carroll © A.L. Carroll Forest health: a closer look One of the most widely used terms in ecosystem management Often in the news Associated with: o o o o Air pollution Invasive species Wildfires Insect outbreaks Powerful personal imagery Connects fragility of “health” with ecosystems Misused and abused… © National Interagency Fire Center Archive, Bugwood.org Healthy or unhealthy? Natural forest: In timber supply area… = pest In protected area… = natural disturbance agent or, if alien invasive,… = pest In multiple use area… = both Interpretation depends on management objectives © A.L. Carroll Healthy or unhealthy? Intensively managed forest: No pest impacts Meets management objectives, but… Limited in all aspects of ecological function © S. Katovich, USDA Forest Service, Bugwood.org Forest health: a functional definition Values create conflicts! Need a definition without human expectations… A working definition: Forest ecosystems are healthy when their underlying ecological processes operate within a natural range of variability, so that on any temporal or spatial scale, they are dynamic and resilient to disturbance How do insects cause disturbance? Insect disturbance is the consequence of herbivory (consumption of plant tissues) Herbivory causes growth loss, deformation, and/or mortality of trees o o Direct impacts (e.g. tissue loss) Indirect impacts (e.g. pathogen transmission) Two major “guilds” of tree herbivores o Foliage feeders (e.g. caterpillars) • Most commonly cause growth loss o © A.L. Carroll Stem feeders (e.g. bark beetles) • Frequently associated with tree mortality © A.L. Carroll Herbivory: eat and grow Herbivory is mainly the job of the juvenile stages There are two distinct types of development by insects: o Complete metamorphosis (holometaboly) • Juvenile stage don’t resemble adults (e.g. larvae) o Incomplete metamorphosis (hemimetaboly) • Juvenile stages do resemble adults (e.g. nymphs) Holometabolous Hemimetabolous Nymph Larva © J.E. Dewey, USDA Forest Service, Bugwood.org Adult Pupa © J.A. Payne, USDA, Bugwood.org Adult © USDA Forest Service, NE Archive, Bugwood.org © USDA Forest Service, R4 Archive, Bugwood.org © D. Riley, Univ. Georgia, Bugwood.org Injury versus damage Injury (i.e. herbivory) o Physical or physiological losses to trees caused by insects (e.g. reduction in leaf area or photosynthesis) Damage o Economic losses caused by insects (e.g. reduction in yield or quality) Feeding Yield loss? (Injury) (Damage) © A.L. Carroll © R.F. Billings, TX Forest Service, Bugwood.org © W.M. Ciesla, Forest Health Mgmt Int, Bugwood.org Injury does not always cause damage! Sucking/galling insects Nature of feeding o Piercing/sucking mouthparts o Plant part specialists o Often prefer healthy/ vigorous hosts o Some species create galls as shelters Nature of injury o Nutrient loss o Pathogen transmission o Phytotoxins and allergic reactions Most serious pests are introduced © W. Cranshaw, CO State Univ., Bugwood.org Sucking/galling insects - examples Insect: Insect: Balsam woolly adelgid (Adelges piceae) Hosts: all true firs Injury: deformation, mortality Cooley spruce gall adelgid (Adelges cooleyi) Hosts: spruces, Douglas-fir Injury: aesthetic (i.e. visual) © L. Livingston, ID Dept. Lands, Bugwood.org © W. Cranshaw, CO State Univ., Bugwood.org © NC Forest Service, Bugwood.org © R.F. Billings, TX Forest Service, Bugwood.org © E.R. Day, VA Polytechnic, Bugwood.org Seed and cone insects Nature of feeding o Direct or indirect feeding on tree reproductive organs o Obligate = feed only on seeds/cones o Facultative = feed on seeds/cones when available Nature of injury o Reduced production of viable seed required for: • Preservation of genetic variation • Response to climate change © L.R. Barber, USDA Forest Service, Bugwood.org Seed and cone insects - examples Insect: Insect: Western conifer seed bug (Leptoglossus occidentalis) Hosts: most conifers Injury: direct seed destruction Fir coneworm (Dioryctria abietivorella) Hosts: most conifers Injury: indirect seed destruction © USDA Forest Service, Odgen Archive, Bugwood.org © L.R. Barber, USDA Forest Service, Bugwood.org © M. Zubrik, Slovakia Forest Res. Inst., Bugwood.org Root and lower stem insects Nature of feeding o Commonly associated with fungi o Root-feeding insects: • Commonly associated with managed stands, increasing importance in forestry o Lower-stem insects: • Commonly associated with mature stands Nature of injury o Consumption of cambial tissues of roots and lower stems o Growth loss, deformation, occasional mortality © G. Csoka, Hungary Forest Res. Inst., Bugwood.org Root and lower-stem insects - examples Insect: Insect: Warren’s root collar weevil (Hylobius warreni) Hosts: lodgepole pine, spruce species Injury: mortality of young trees Red turpentine beetle (Dendroctonus valens) Hosts: lodgepole pine, ponderosa pine Injury: mortality of stressed trees © Joseph Berger, Bugwood.org © A.L. Carroll © C. Hellqvist, Swedish Univ. Agric. Sci., Bugwood.org © E.H. Holsten, USDA Forest Service, Bugwood.org © A.L. Carroll © USDA Forest Service, NE Area Archive, Bugwood.org Shoot insects Nature of feeding o Specialization on elongating shoots o Preference for young, vigorous trees Nature of injury o Growth loss, longer rotation o Malformation o Impacts greatest on young trees o Indirect mortality - competition o Injury often positively related to tree vigour, increased by fertilization, irrigation, spacing © E.B. Walker, VT Dept. Forests Parks & Rec., Bugwood.org Shoot insects - examples Insect: Insect: White pine weevil (spruce weevil) (Pissodes strobi) Hosts: spruce and pine species Injury: growth loss, deformation Bud moths (Zeiraphera spp.) Hosts: spruce and larch species Injury: growth loss, deformation © A.L. Carroll © S. Katovich, USDA Forest Service, Bugwood.org © A.S. Munson, USDA Forest Service, Bugwood.org © W.M. Ciesla, Forest Health Mgmt Int, Bugwood.org © CT Agric. Exp. Stn. Archive, Bugwood.org © A.L. Carroll Woodborers Nature of feeding o Develop in phloem and sapwood o Associated with stressed, dead trees o Attracted to volatiles associated with wood deterioration Nature of injury o Kill or weaken live trees (typically stressed) o Some species associated with fungi o Impact wood products o Significant potential as invasive species © D. Cappaert, Michigan State Univ., Bugwood.org Wood borers - examples Insect: Insect: Golden buprestid (Buprestis aurulenta) Hosts: Douglas-fir, pine species Injury: sapwood boring (reduced integrity) Striped ambrosia beetle (Trypodendron lineatum) Hosts: most conifers Injury: sapwood boring (aesthetics) © M. Jurc, Univ. Ljubljana, Bugwood.org © S. Tunnock, USDA Forest Service, Bugwood.org © A.L. Carroll © M. Jurc, Univ. Ljubljana, Bugwood.org © P. Kapitola, Czechia State Phytosan. Admin., Bugwood.org Defoliators Nature of feeding o General preference for young foliage o Some species feed on old foliage o Synchrony between egg hatch and bud burst usually required Nature of injury o Feeding causes loss of photosynthetic organs o General loss of tree vigour o Growth reduction, crown deformation o Occasional “top-kill” or mortality o Several species prone to outbreaks © A.L. Carroll Defoliators - examples Insect: Insect: Western spruce budworm (Choristoneura occidentalis) Hosts: Douglas-fir, other conifers Injury: growth loss, mortality Douglas-fir tussock moth (Orgyia pseudotsugata) Hosts: Douglas-fir Injury: growth loss, mortality © K.E. Gibson, USDA Forest Service, Bugwood.org © D. Owen, CA Dept. Forestry and Fire Prot., Bugwood.org © S. Tunnock, USDA Forest Service, Bugwood.org © W.M. Ciesla, Forest Health Mgmt Int, Bugwood.org © D. McComb, USDA Forest Service, Bugwood.org Bark beetles Nature of feeding o General preference for older trees o Feed beneath bark on phloem tissues o Complex adaptations to overcome tree defenses Nature of injury o For many species, successful reproduction requires tree death o Close association with pathogenic fungi o Several species prone to outbreaks © A.L. Carroll Bark beetle impacts in recent decades Bark beetles - examples Insect: Insect: Mountain pine beetle (Dendroctonus ponderosae) Hosts: pine species Damage: tree mortality Spruce beetle (Dendroctonus rufipennis) Hosts: spruce species Damage: tree mortality © E.H. Holsten, USDA Forest Service, Bugwood.org © A.L. Carroll © A.L. Carroll © E.H. Holsten, USDA Forest Service, Bugwood.org © A.L. Carroll © W.M. Ciesla, Forest Health Mgmt Int, Bugwood.org Some final words… Insects can inflict injury to all parts of a tree Disturbance arising from injury extends across spatial scales – from trees to landscapes Insect disturbance is a critical component of forest health Most insect herbivory is natural, some is associated with introduced species o Management objective and actions must vary accordingly Thank you! © A.L. Carroll