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Predators of The^ Spruce Budworm I Jí^íS. United States fil X m Department of Spruce m Agriculture Budworms Forest Service Handbook Cooperative State Research Service Agriculture Handbook No. 644 Predators of the^ Spruce Budworm i 864305 In 1977, the United States Depart- strategies for controlling the spruce ment of Agriculture and the Canadian budworms and managing budworm- Department of the Environment susceptible forests to help forest agreed to cooperate in an expanded managers attain their objectives in an and accelerated research and develop- economically and environmentally ment effort, the Canada/United States acceptable manner. The work Spruce Budworms Program (C ANUS A), represented in this pubUcation was aimed at the spruce budworm in the wholly or partially funded by the East and the western spruce budworm Program. This manual is one in a in the West. The objective of series on the spruce budworm. CANUSA was to design and evaluate Canada United States Spruce Budworms Program November 1985 Contents Introduction 1 Life History of the Spruce Budworm 2 Host Trees 2 Susceptibility of Life Stages to Prédation 3 Predator Groups 5 Overview of Predators 5 Predators of Eggs 6 Phalangids 6 Mites 6 Spiders 8 Plant Bugs 10 Lacewings 10 Beetles 10 Ants 11 Birds 12 Predators of Small Larvae 13 Phalangids 13 Spiders 14 Beetles 17 Ants 17 Miscellaneous Insects 18 Birds 18 Predators of Large Larvae 19 Spiders 19 Dragonflies 20 Beetles 21 Spruce Coneworms 24 Ants 25 Wasps 27 Fish 30 Mammals 30 Birds 31 Predators of Pupae 34 Spiders 34 Beetles 34 Spruce Coneworms 35 Budworm Larvae 37 Syrphids 37 Ants 37 Mammals 38 Birds 38 Predators of Adults 41 Spiders 41 Dragonflies 42 Beetles 45 Robber Flies 45 Ants 46 Birds 47 Predators as Natural Regulators 48 Endemic Spruce Budworm Populations 49 Epidemic Spruce Budworm Populations 52 Methods for Studying Prédation 56 Exclusion Techniques 56 Direct Assessments 57 Serological Tests 57 Radioisotope Tagging 57 Direct Observations 58 Conservation and Enhancement of Predators 59 Habitat Requirements 59 Food Requirements 59 Nesting Requirements 60 Summary 61 Acknowledgments 62 Selected References and Guides for Identifying Predators 62 Phalangids 62 Spiders 62 Mites 63 Dragonflies 63 Carabid Beetles 63 Coneworms 63 Robber Flies 64 Ants 64 Wasps 64 Fish 64 Birds 64 Manmials 64 Literature Cited 65 Predators of the Spruce Budworm by Daniel T. Jennings and Hewlette S. Crawford, Jr.^ Introduction The spruce budworm, Choristoneura outbreaks. These natural agents of fumiferana (Clemens) (Lepidoptera: control, also called natural enemies, Tortricidae), is the most destructive include various diseases, parasites, forest insect pest in eastern North and predators. America. Millions of acres of spruce-fir forests have been damaged Enhancement of predators of the by the spruce budworm in eastern spruce budworm is a desirable feature Canada and in the North-central and of integrated pest management; Northeastern United States from Min- however, before we can incorporate nesota to Maine. Recorded outbreaks predator enhancement we need to of the budworm date back to the know which predators are important, early 1700's; but in the 20th century, how effective they are in regulating outbreaks are increasing in frequency, budworm populations, and what their extent, and severity (Biais 1983). habitat requirements are. Ultimately, such information and understanding The spruce budworm is a native, will lead to the identification and natural component of the spruce-fir development of forest-management forest. Managers need to know how strategies that enhance predators of to manage the insect to prevent or the spruce budworm. minimize damage to the forest. Applied control of the spruce bud- The goals of this handbook are to worm has been largely chemical. A (1) review available information on more integrated approach is needed predators of the spruce budworm; that includes safer, longer lasting, (2) provide information on predator less costly methods of pest manage- biologies, including life stages, modes ment. Such methods should be com- of attack, and general importance; patible with multiple uses of all forest (3) summarize the importance of resources, including fiber, wildlife predators and prédation in population and fish production, soil and water dynamics of the spruce budworm; conservation, and recreation. (4) review forest-management prac- tices that enhance predator popula- The spruce-fir forest has various tions; and (5) list basic literature natural agents that help keep bud- sources for identifying predators of worm populations in check between the spruce budworm. 'Daniel Jennings is a research forest ento- mologist with the USD A Forest Service, North- eastern Forest Experiment Station, Orono, ME. Hewlette Crawford, Jr., is a research wildlife biologist with the Station at Amherst, MA. Life History of the Spruce Budworm The spruce budworm has four life two or more shoots together to form stages: egg, larva, pupa, and adult. a feeding tunnel or shelter. During The adult stage is a moth with a this active feeding period, the larvae wingspan of about three-quarters of molt three more times to reach the an inch (2 cm). The female moths lay sixth instar by late June or early July. their eggs in masses on needles of The fifth and sixth instars cause most host trees in mid-July to early of the feeding damage to host trees. August. The number of eggs per Destruction of current shoots causes mass varies considerably but averages considerable larval movement, and about 20. The eggs hatch in about 8 many larvae drop from their host to 12 days, and the newly emerged trees and are exposed to ground- first-instar larvae do not feed but inhabiting predators. disperse and seek overwintering sites under bark scales and in old After development, the larva stops staminate flower bracts. After spin- feeding and transforms into a pupa. ning a silken hibernaculum, the larva The pupal stage lasts about 10 days, molts to the second instar, enters after which the moth emerges. The diapause, and overwinters. female moth emits a sex pheromone that attracts the male for mating. In the spring of the following year, Moths live about 2 weeks and are the second-instar larvae emerge from found from early July to August. their hibernacula, disperse, and seek Shortly after mating, the female moth feeding sites. Newly opened staminate begins to lay eggs, completing the flowers are the preferred food on life cycle. There is only one genera- balsam fir, Abies balsamea (L.) tion of the spruce budworm per year, Miller, during the initial feeding but the life cycle spans 2 calendar stage. When staminate flowers are years. not available and vegetative buds are "tight" and unexpanded, larvae mine Host Trees the previous year's needles. Usually, only one needle is mined and the Balsam fir is the principal host of the larva molts to the third instar within spruce budworm in eastern North or soon after leaving the needle. By America; it is the host that suffers the mid- to late May or early June, greatest damage and tree mortality depending on temperatures, most of (Miller 1963a). Outbreaks are most the larvae leave mined needles and likely when mature stands of balsam begin feeding on newly opened fir cover extensive areas (Biais 1983). vegetative buds (Miller 1963a). The budworm also attacks and feeds on white spruce, Picea glauca Feeding continues on the new foliage (Moench) Voss; red spruce, of developing shoots and on old P. rubens Sarg.; black spruce, P. foliage once the current foliage is mariana (Mill.) B.S.P.; and blue consumed. The larvae typically web spruce, P. pungens Engelm. During outbreaks there may be feeding on to the forest floor, where ants, tamarack, Larix lancina (Du Roi) K. carabid beetles, spiders, and small Koch; eastern white pine, Pinus mammals often are numerous. strobus L.; and eastern hemlock, Tsuga canadensis (L.) Carr. Mature Conversely, the inability to move and and overmature trees are most escape predators increases the suscep- susceptible to attack, though trees of tibility of pupae to prédation. Pupae all sizes can be damaged or killed generally are found near the apexes during an epidemic. of branches in silken shelters con- structed by the larvae before pupa- Susceptibility of Life Stages tion. Although capable of wiggling to Prédation and squirming, pupae usually remain attached to these silken threads by All life stages of the spruce budworm means of cremaster hooks. are susceptible to attack by predators. However, some stages are more sus- Spruce budworm moths may escape ceptible than others. For example, some predators by taking flight; during both larval dispersal periods, however, flying moths are exposed to the small larvae are exposed and sub- aerial-searching predators such as ject to attack by arboreal and epigeal dragonflies, robber flies, and birds. predators. Once needle or bud mining There are temporal and sexual dif- begins, the larvae are protected and ferences in susceptibilities of moths to are less likely to be eaten by pred- prédation. Male spruce budworm ators. Overwintering larvae in hiber- moths are more active, rapid fliers nacula are protected but not immune than female moths. Female moths to prédation. must lay some of their eggs before engaging in long-distance flights. Many predators rely on movements to Moths are subject to prédation by detect potential prey. Because egg foliage-searching predators such as masses are deposited on needles and birds and spiders. During coitus, both are relatively stable, the egg masses sexes are susceptible and vulnerable are not likely to be detected except to attack because they represent a by foliage-searching predators such as relatively large target and their coccinellid beetles and birds. movements are somewhat impaired. Large larvae of the spruce budworm Habitats occupied by the different life may escape foliage-searching stages of the spruce budworm and predators by dropping from their host searching behaviors of potential trees.
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