Skip to main content Skip to main navigation menu Skip to site footer
Type: Articles
Published: 2005-04-12
Page range: 1–46
Abstract views: 122
PDF downloaded: 29

A review of the tachinid parasitoids (Diptera: Tachinidae) of Nearctic Choristoneura species (Lepidoptera: Tortricidae), with keys to adults and puparia

Invertebrate Biodiversity, Agriculture and Agri-Food Canada, 960 Carling Avenue, Ottawa, Ontario, Canada, K1A 0C6
Diptera Tachinidae Choristoneura Nearctic keys

Abstract

The genus Choristoneura (Lepidoptera: Tortricidae) comprises about 16 species in the Nearctic Region and includes several destructive pests of importance to forestry and agriculture. The following 15 species of Tachinidae (Diptera) are recognized as occasional or common parasitoids of Choristoneura species in this region: Actia diffidens Curran, Actia interrupta Curran, Ceromasia auricaudata Townsend, Compsilura concinnata (Meigen), Cyzenis incrassata (Smith), Eumea caesar (Aldrich), Hemisturmia parva (Bigot), Hyphantrophaga blanda (Osten Sacken), Hyphantrophaga virilis (Aldrich and Webber), Lypha fumipennis Brooks, Madremyia saundersii (Williston), Nemorilla pyste (Walker), Nilea erecta (Coquillett), Phryxe pecosensis (Townsend), and Smidtia fumiferanae (Tothill). Keys to the adults and puparia of these tachinid species are provided. The known distribution, biology, and rates of parasitism are reviewed for each species and published host records from Choristoneura species are listed. Colour habitus images of adults, illustrations of all puparia, and illustrations of features mentioned in the adult key are included. Thirteen other species of Tachinidae recorded from Choristoneura species but excluded from the main text as highly dubious, rare, or accidental records are briefly discussed.

References

  1. Aldrich, J.M. (1916) Two new Canadian Diptera. Canadian Entomologist, 48, 20–22.

    Aldrich, J.M. & Webber, R.T. (1924) The North American species of parasitic two-winged flies belonging to the genus Phorocera and related genera. Proceedings of the United States National Museum, 63 (Art. 17) [= No. 2486], 1–90.

    Allen, D.C., Knight, F.B., Foltz, J.L. & Mattson, W.J. (1969) Influence of parasites on two populations of Choristoneura pinus (Lepidoptera: Tortricidae) in Michigan. Annals of the Entomological Society of America, 62, 1469–1475.

    Arnaud, P.H., Jr. (1978) A host-parasite catalog of North American Tachinidae (Diptera). United States Department of Agriculture, Miscellaneous Publication, 1319, 860 pp.

    Bedard, W.D. (1938) An annotated list of the insect fauna of Douglas fir (Pseudotsuga mucronata rafinesque) in the northern Rocky Mountain region. Canadian Entomologist, 70, 188–197.

    Benjamin, D.M. & Drooz, A.T. (1954) Parasites affecting the jack-pine budworm in Michigan. Journal of Economic Entomology, 47, 588–591.

    Bigot, J.-M.-F. (1889) Diptères nouveaux ou peu connus. 34e partie. XLII. Diagnoses de nouvelles espèces. Annales de la Société Entomologique de France, Série 6, 8, 253–270.

    Blais, J.R. (1960) Spruce budworm parasite investigations in the Lower St. Lawrence and Gaspé regions of Quebec. Canadian Entomologist, 92, 384–396.

    Blais, J.R. (1965) Parasite studies in two residual spruce budworm (Choristoneura fumiferana (Clem.)) outbreaks in Quebec. Canadian Entomologist, 97, 129–136.

    Boettner, G.H., Elkinton, J.S. & Boettner, C.J. (2000) Effects of a biological control introduction on three nontarget native species of saturniid moths. Conservation Biology, 14, 1798–1806.

    Bourchier, R.S. (1991) Growth and development of Compsilura concinnata (Meigan [sic]) (Diptera: Tachinidae) parasitizing gypsy moth larvae feeding on tannin diets. Canadian Entomologist, 123, 1047–1055.

    Bourchier, R.S. & Smith, S.M. (1998) Interactions between large-scale inundative releases of Trichogramma minutum (Hymenoptera: Trichogrammatidae) and naturally occurring spruce budworm (Lepidoptera: Tortricidae) parasitoids. Environmental Entomology, 27, 1273–1279.

    Brooks, A.R. (1945) New Canadian Diptera (Tachinidae). Canadian Entomologist, 77, 78–96.

    Brown, A.W.A. (1941) Foliage insects of spruce in Canada. Canada Department of Agriculture Publication, 712 (Technical Bulletin 31), 1–29.

    Burgess, A.F. & Crossman, S.S. (1927) The satin moth, a recently introduced pest. United States Department of Agriculture, Department Bulletin, 1469, 1–22 + 1 pl.

    Caesar, L. (1916) Leaf-rollers attacking apples. Annual Report of the Entomological Society of Ontario [for] 1915, 46, 163–178.

    Cappuccino, N., Houle, M.-J. & Stein, J. (1999) The influence of understory nectar resources on parasitism of the spruce budworm Choristoneura fumiferana in the field. Agricultural and Forest Entomology, 1, 33–36.

    Cappuccino, N., Lavertu, D., Bergeron, Y. & Régnière, J. (1998) Spruce budworm impact, abundance and parasitism rate in a patchy landscape. Oecologia, 114, 236–242.

    Carolin, V.M. & Coulter, W.K. (1959) The occurrence of insect parasites of Choristoneura fumiferana (Clem.), in Oregon. Journal of Economic Entomology, 52, 550–555.

    Ciesla, W.M. (1964) Life history and habits of the elm spanworm, Ennomos sibsignarius, in the southern Appalachian Mountains (Lepidoptera: Geometridae). Annals of the Entomological Society of America, 57, 591–596.

    Clausen, C.P. (1978) Tortricidae. In: Clausen, C.P. (Ed.), Introduced parasites and predators of arthropod pests and weeds: a world review. United States Department of Agriculture, Agriculture Handbook, 480, 236–240.

    Coppel, H.C. (1947) The collection of spruce budworm parasites in British Columbia with notes on their overwintering habits. Annual Report of the Entomological Society of Ontario [for] 1946, 77, 38–40.

    Coppel, H.C. (1953) Effects of super, — multi, — and hyperparasitism on the abundance of Phytodietus fumiferanae Rohw. (Hymenoptera: Ichneumonidae), a primary parasite of the spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae). Report of the Quebec Society for the Protection of Plants [for] 1950–1951, 32–33, 189–193.

    Coppel, H.C. (1958) Studies on dipterous parasites of the spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae). VI. Phorocera incrassata Smith (Diptera: Tachinidae). Canadian Journal of Zoology, 36, 453–462.

    Coppel, H.C. (1960) Key to adults of dipterous parasites of spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae). Annals of the Entomological Society of America, 53, 94–97.

    Coppel, H.C. & Maw, M.G. (1954a) Studies on dipterous parasites of the spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae). III. Ceromasia auricaudata Tns. (Diptera: Tachinidae). Canadian Journal of Zoology, 32, 144–156.

    Coppel, H.C. & Maw, M.G. (1954b) Studies on dipterous parasites of the spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae). IV. Madremyia saundersii (Will.) (Diptera: Tachinidae). Canadian Journal of Zoology, 32, 314–323.

    Coppel, H.C. & Smith, B.C. (1957) Studies on dipterous parasites of the spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae). V. Omotoma fumiferanae (Tot.) (Diptera: Tachinidae). Canadian Journal of Zoology, 35, 581–592.

    Coquillett, D.W. (1902) New Diptera from North America. Proceedings of the United States National Museum, 25 [= No. 1280], 83–126.

    Cossentine, J., Jensen, L., Deglow, E., Bennett, A., Goulet, H., Huber, J. & O’Hara, J. (2004) The parasitoid complex affecting Choristoneura rosaceana and Pandemis limitata in organically managed apple orchards. BioControl, 49, 359-372.

    Culver, J.J. (1919) A study of Compsilura concinnata, an imported tachinid parasite of the gipsy moth and the brown-tail moth. United States Department of Agriculture, Bureau of Entomology, Bulletin, 766, 1–27 + 1 pl.

    Curran, C.H. (1933) The North American species of Actia in the American Museum of Natural History. American Museum Novitates, 614, 1–7.

    Cusson, M., Laforge, M., Régnière, J., Béliveau, C., Trudel, D., Thireau, J.C., Bellemare, G., Keirstead, N. & Stolz, D. (2002) Multiparasitism of Choristoneura fumiferana by the ichneumonid Tranosema rostrale and the tachinid Actia interrupta: occurrence in the field and outcome of competition under laboratory conditions. Entomologia Experimentalis et Applicata, 102, 125–133.

    Dang, P.T. (1992) Morphological study of male genitalia with phylogenetic inference of Choristoneura Lederer (Lepidoptera: Tortricidae). Canadian Entomologist, 124, 7–48.

    Daviault, L. (1946) Forest entomology. Report of the Minister of Lands and Forests of the province of Quebec for the year ending March 31st, 1945, 113–122.

    Daviault, L. (1950) Les parasites de la tordeuse des bourgeons de l’épinette (Choristoneura fumiferana Clem.) dans la province de Québec. Report of the Quebec Society for the Protection of Plants, 31, 41–45.

    Davis, J.J. (1912) Report on insects injurious to flowering and ornamental greenhouse plants in Illinois. Report of the State Entomologist on the noxious and beneficial insects of the state of Illinois, 27, 83–143.

    Dixon, J.C. & Benjamin, D.M. (1963) Natural control factors associated with the jack-pine budworm, Choristoneura pinus. Journal of Economic Entomology, 56, 266–270.

    Doganlar, M. & Beirne, B.P. (1978) Natural enemies of budworms, Choristoneura spp. (Lepidoptera: Tortricidae), on Douglas fir near Yale, British Columbia, in 1977. Journal of the Entomological Society of British Columbia, 75, 25–26.

    Dowden, P.B., Buchanan, W.D. & Carolin, V.M. (1948) Natural-control factors affecting the spruce budworm. Journal of Economic Entomology, 41, 457–464.

    Dowden, P.B., Carolin, V.M. & Dirks, C.O. (1951) Natural control factors affecting the spruce budworm in the Adirondacks during 1946–1948. Journal of Economic Entomology, 43 (1950), 774–783.

    Dowden, P.B., Jaynes, H.A. & Carolin, V.M. (1953) The role of birds in a spruce budworm outbreak in Maine. Journal of Economic Entomology, 46, 307–312.

    Drooz, A.T. & Benjamin, D.M. (1956) Parasites from two jack-pine budworm outbreaks on the Upper Peninsula of Michigan. Journal of Economic Entomology, 49, 412–413.

    Evans, D. (1962) Descriptions and life history of Melanolophia imitata (Walker) (Lepidoptera: Geometridae). Canadian Entomologist, 94, 594–605.

    Fitzpatrick, S.M., Troubridge, J.T. & Maurice, C. (1994) Parasitoids of blackheaded fireworm (Rhopobota naevana Hbn.) larvae on cranberries, and larval escape behaviour. Journal of the Entomological Society of British Columbia, 91, 73–74.

    Frankenhuyzen, K. van. (2002) Choristoneura pinus pinus Freeman, jack pine budworm (Lepidoptera: Tortricidae). In: Mason, P.G. & Huber, J.T. (Eds.), Biological control programmes in Canada, 1981–2000. CABI Publishing, Wallingford and New York, pp. 75–78.

    Franklin, H.J. (1943) Injurious and beneficial insects affecting the cranberry. Massachusetts Agricultural Experiment Station Bulletin, 398, 26–28.

    Franklin, H.J. (1950) Cranberry insects in Massachusetts. Part V. Insects and other animals beneficial in cranberry growing. Massachusetts Agricultural Experiment Station Bulletin, 445, 55–76.

    Gardiner, J.G. (1946) Report of the Minister of Agriculture [for] 1945–46. Report of the Minister of Agriculture for the Dominion of Canada for the year ended March 31, 1946, 235 pp.

    Graham, A.R. (1965) A preliminary list of the natural enemies of Canadian agricultural pests. Canada Department of Agriculture Research Institute Information Bulletin, 4, 1–179.

    Graham, A.R. & Jones, P.A. (1962) Appendix I. Parasites and predators released against forest insects in Canada, 1910–58. In: McLeod, J.H., McGugan, B.M. & Coppel, H.C., A review of the biological control attempts against insects and weeds in Canada. Commonwealth Institute of Biological Control, Technical Communication, 2, 129–175.

    Hagley, E.A.C. & Barber, D.R. (1991) Foliage-feeding Lepidoptera and their parasites recovered from unmanaged apple orchards in southern Ontario. Proceedings of the Entomological Society of Ontario, 122, 1–7.

    Harris, J.W.E. & Dawson, A.F. (1979) Parasitoids of the western spruce budworm, Choristoneura occidentalis (Lepidoptera: Tortricidae), in British Columbia 1977–78. Journal of the Entomological Society of British Columbia, 76, 30–38.

    Harris, J.W.E. & Dawson, A.F. (1985) Parasitoids of the western spruce budworm in British Columbia. In: Sanders, C.J., Stark, R.W., Mullins, E.J. & Murphy, J. (Eds.), Recent advances in spruce budworms research. Proceedings of the CANUSA spruce budworms research symposium. Bangor, Maine, September 16–20, 1984. Canadian Forestry Service, Ottawa, p. 102.

    Hébert, C. & Cloutier, C. (1990a) Temperature-dependent development of eggs and larvae of Winthemia fumiferanae Toth. (Diptera: Tachinidae), a larval-pupal parasitoid of the spruce budworm (Lepidoptera: Tortricidae). Canadian Entomologist, 122, 329–341.

    Hébert, C. & Cloutier, C. (1990b) Host instar as a determinant of preference and suitability for two parasitoids attacking late instars of the spruce budworm (Lepidoptera: Tortricidae). Annals of the Entomological Society of America, 83, 734–741.

    Hébert, C., Cloutier, C. & Régnière, J. (1990) Factors affecting the flight activity of Winthemia fumiferanae (Diptera: Tachinidae). Environmental Entomology, 19, 293–302.

    Hébert, C., Cloutier, C., Régnière, J. & Perry, D.F. (1989) Seasonal biology of Winthemia fumiferanae Toth. (Diptera: Tachinidae), a larval-pupal parasitoid of the spruce budworm (Lepidoptera: Tortricidae). Canadian Journal of Zoology, 67, 2384–2391.

    Herting, B. (1984) Catalogue of Palearctic Tachinidae (Diptera). Stuttgarter Beiträge zur Naturkunde Serie A (Biologie), 369, 1–228.

    Hewitt, C.G. (1913) Report of the Dominion Entomologist for the year ending March 31 1912. Canada Department of Agriculture, Annual Report on Experimental Farms for the year 1911–12, 173–189.

    Huber, J.T., Eveleigh, E., Pollock, S. & McCarthy, P. (1996) The chalcidoid parasitoids and hyperparasitoids (Hymenoptera: Chalcidoidea) of Choristoneura species (Lepidoptera: Tortricidae) in America north of Mexico. Canadian Entomologist, 128, 1167–1220.

    Ichiki, R. & Shima, H. (2003) Immature life of Compsilura concinnata (Meigen) (Diptera: Tachinidae). Annals of the Entomological Society of America, 96, 161–167.

    Jaynes, H.A. & Drooz, A.T. (1952) The importance of parasites in the spruce budworm infestations in New York and Maine. Journal of Economic Entomology, 45, 1057–1061.

    Johannsen, O.A. (1913) Spruce bud worm and spruce leaf miners. Maine Agricultural Experiment Station, Bulletin, 210, 13–36 + 3 pls.

    Johnson, C.W. (1925) Fauna of New England. 15. List of the Diptera or two-winged flies. Occasional papers of the Boston Society of Natural History, 7, 1–326.

    Knowlton, G.F. & Allen, M.W. (1937) Oblique-banded leaf roller, a dewberry pest in Utah. Journal of Economic Entomology, 30, 780–785.

    Knowlton, G.F. & Hansen, L.L. (1938) Notes on berry insects of Utah. Proceedings of the Utah Academy of Sciences, Arts and Letters, 15, 127–130.

    Kulman, H.M. & Hodson, A.C. (1961) Parasites of the jack-pine budworm, Choristoneura pinus, with special reference to parasitism at particular stand locations. Journal of Economic Entomology, 54, 221–224.

    Li, S.Y., Fitzpatrick, S.M., Hueppelsheuser, T., Cossentine, J.E. & Vincent, C. (2002) Choristoneura rosaceana (Harris), obliquebanded leafroller (Lepidoptera: Tortricidae). In: Mason, P.G. & Huber, J.T. (Eds.), Biological control programmes in Canada, 1981–2000. CABI Publishing, Wallingford and New York, pp. 78–83.

    Li, S.Y., Fitzpatrick, S.M., Troubridge, J.T., Sharkey, M.J., Barron, J.R. & O’Hara, J.E. (1999) Parasitoids reared from the obliquebanded leafroller (Lepidoptera: Tortricidae) infesting raspberries. Canadian Entomologist, 131, 399–404.

    MacDonald, D.R. & Webb, F.E. (1963) Insecticides and the spruce budworm. In: Morris, R.F. (Ed.), The dynamics of epidemic spruce budworm populations. Memoirs of the Entomological Society of Canada, 31, 288–310 + figs. 39.1–39.6.

    Maw, M.G. & Coppel, H.C. (1953) Studies on dipterous parasites of the spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae). II. Phryxe pecosensis (Tns.) (Diptera: Tachinidae). Canadian Journal of Zoology, 31, 392–403.

    McAlpine, J.F. (1981) Morphology and terminology – adults. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Volume 1. Agriculture Canada Monograph, 27, 9–63.

    McGugan, B.M. & Blais, J.R. (1959) Spruce budworm parasite studies in northwestern Ontario. Canadian Entomologist, 91, 758–783.

    McGugan, B.M. & Coppel, H.C. (1962) Part II — Biological control of forest insects, 1910–1958. In: McLeod, J.H., McGugan, B.M. & Coppel, H.C., A review of the biological control attempts against insects and weeds in Canada. Commonwealth Institute of Biological Control, Technical Communication, 2, 35–127.

    McKnight, M.E. (1974) Parasitoids of the western spruce budworm in Colorado. Environmental Entomology, 3, 186–187.

    Meigen, J.W. (1824) Systematische Beschreibung der bekannten europäischen zweiflügeligen Insekten. Hamm, Vol. 4, xii + 428 pp.

    Mellini, E. (1990) Sinossi di biologia dei Ditteri Larvevoridi. Bollettino dell’Istituto di Entomologia “Guido Grandi” dell’Università di Bologna, 45, 1–38.

    Miller, C.A. (1955) A technique for assessing spruce budworm larval mortality caused by parasites. Canadian Journal of Zoology, 33, 5–17.

    Miller, C.A. (1963) Parasites and the spruce budworm. In: Morris, R.F. (Ed.), The dynamics of epidemic spruce budworm populations. Memoirs of the Entomological Society of Canada, 31, 228–244 + figs. 34.1–34.7.

    Nealis, V.G. (1991) Parasitism in sustained and collapsing populations of the jack pine budworm, Choristoneura pinus pinus Free. (Lepidoptera: Tortricidae), in Ontario, 1985–1987. Canadian Entomologist, 123, 1065–1076.

    Neunzig, H.H. & Gyrisco, G.G. (1955) Some insects injurious to birdsfoot trefoil in New York. Journal of Economic Entomology, 48, 447–450.

    O’Hara, J.E. (1985) Oviposition strategies in the Tachinidae, a family of beneficial flies. University of Alberta, Agriculture and Forestry Bulletin, 8(2), 31–34.

    O’Hara, J.E. (1988) Survey of first instars of the Siphonini (Diptera: Tachinidae). Entomologica Scandinavica, 18, 367–382.

    O’Hara, J.E. (1991) Revision of Nearctic species of Actia Robineau-Desvoidy (Diptera: Tachinidae). Canadian Entomologist, 123, 745–776.

    O’Hara, J.E. (2002) Revision of the Polideini (Tachinidae) of America north of Mexico. Studia dipterologia, Supplement, 10, 170 pp.

    O’Hara, J.E. & Cooper, B.E. (1992) Revision of the Nearctic species of Cyzenis Robineau-Desvoidy (Diptera: Tachinidae). Canadian Entomologist, 124, 785–813.

    O’Hara, J.E. & Wood, D.M. (2004) Catalogue of the Tachinidae (Diptera) of America north of Mexico. Memoirs on Entomology, International, 18, 410 pp.

    Osten Sacken, C.R. (1887) Some North American Tachinae. Canadian Entomologist, 19, 161–166.

    Otvos, I.S. (1991) Tortricids in forestry. North American species in forestry. In: van der Geest, L.P.S. & Evenhuis, H.H. (Eds.), Tortricid pests. Their biology, natural enemies and control. World Crop Pests 5, Elsevier, Amsterdam, pp. 719–756.

    Pfannenstiel, R.S. & Unruh, T.R. (2003) Conservation of leafroller parasitoids through provision of alternate hosts in near-orchard habitats. In: Van Driesche, R.G. (Ed.), Proceedings of the International Symposium on Biological Control of Arthropods. Honolulu, Hawaii, 14–18 January 2002. United States Department of Agriculture, Forest Service, Morgantown, WV. FHTET-2003-05, pp. 256–262.

    Prebble, M.L. (1935) Actia diffidens Curran, a parasite of Peronea variana (Fernald) in Cape Breton, Nova Scotia. Canadian Journal of Research, 12, 216–227.

    Pree, D.J., Whitty, K.J., Pogoda, M.K. & Bittner, L.A. (2001) Occurrence of resistance to insecticides in populations of the obliquebanded leafroller from orchards. Canadian Entomologist, 133, 93–103.

    Prentice, R.M. (1955) The life history and some aspects of the ecology of the large aspen tortrix, Choristoneura conflictana (Wlkr.) (n. comb.) (Lepidoptera: Tortricidae). Canadian Entomologist, 87, 461–473.

    Raizenne, H. (1952) Forest Lepidoptera of southern Ontario and their parasites received and reared at the Ottawa Forest Insect Survey Laboratory from 1937 to 1948. Canada Department of Agriculture, Science Service, Division of Forest Biology, Ottawa, v + 277 pp.

    Reeks, W.A., Forbes, R.S. & Cuming, F.G. (1948) Maritime provinces and Newfoundland. Canada Department of Agriculture, Science Service, Annual Report of the Forest Insect Survey, Forest Insect Investigations, 1947, 9–23.

    Richmond, H.A. (1938) Forest insects of the season 1937 in Manitoba and northern Ontario. Canadian Insect Pest Review, 16, 45–46.

    Richmond, H.A. (1940) Review of forest insects of the season 1939 from the Winnipeg laboratory. Canadian Insect Pest Review, 18, 49–54.

    Richmond, H.A. (1941a) Forest insects of the season 1940 in central Canada. Canadian Insect Pest Review, 19, 57–60.

    Richmond, H.A. (1941b) [Untitled.] Canadian Insect Pest Review, 19, 209.

    Roland, J., Evans, W.G. & Myers, J.H. (1989) Manipulation of oviposition patterns of the parasitoid Cyzenis albicans (Tachinidae) in the field using plant extracts. Journal of Insect Behavior, 2, 487–503.

    Ross, D.A. (1952) Key to the puparia of the dipterous parasites of Choristoneura fumiferana Clem. Canadian Entomologist, 84, 108–112.

    Royama, T. (1984) Population dynamics of the spruce budworm Choristoneura fumiferana. Ecological Monographs, 54, 429–462.

    Sabrosky, C.W. & Arnaud, P.H., Jr. (1965) Family Tachinidae (Larvaevoridae). In: Stone, A., Sabrosky, C.W., Wirth, W.W., Foote, R.H. & Coulson, J.R. (Eds.), A catalog of the Diptera of America north of Mexico. United States Department of Agriculture, Agriculture Handbook, 276, 961–1108.

    Sanders, C.J. (1991) Tortricids in forestry. Biology of North American spruce budworms. In van der Geest, L.P.S. & Evenhuis, H.H. (Eds.), Tortricid pests. Their biology, natural enemies and control. World Crop Pests 5, Elsevier, Amsterdam, pp. 579–620.

    Schaffner, J.V., Jr. (1959) Microlepidoptera and their parasites reared from field collections in the northeastern United States. United States Department of Agriculture, Miscellaneous Publication, 767, iv + 97 pp.

    Schaffner, J.V., Jr. & Griswold, C.L. (1934) Macrolepidoptera and their parasites reared from field collections in the northeastern part of the United States. United States Department of Agriculture, Miscellaneous Publication, 188, 1-160.

    Schaupp, W.C. Jr., Volney, W.J.A. & Waters, W.E. (1991) Parasitoids of endemic and epidemic populations of Choristoneura occidentalis Freeman and Choristoneura retiniana (Walsingham) (Lepidoptera: Tortricidae) in southern Oregon. Canadian Entomologist, 123, 1095–1102.

    Schmid, J.M. (1981) Distribution of western spruce budworm (Lepidoptera: Tortricidae) insect parasites in the crowns of host trees. Canadian Entomologist, 113, 1101–1106.

    Schoenmaker, A., Cusson, M. & Frankenhuyzen, K. van. (2001) Interactions between Bacillus thuringiensis and parasitoids of late-instar larvae of the spruce budworm (Lepidoptera: Tortricidae). Canadian Journal of Zoology, 79, 1697–1703.

    Schröder, D. (1969) Lypha dubia (Fall.) (Dip.: Tachinidae) as a parasite of the European pine shoot moth, Rhyacionia buoliana (Schiff.) (Lep.: Eucosmidae) in Europe. Commonwealth Institute of Biological Control, Technical Bulletin, 12, 43–60.

    Schuh, J. & Mote, D.C. (1948) The oblique-banded leaf roller on red raspberries [Archips rosaceana (Harris)]. Oregon Agricultural Experiment Station, Technical Bulletin, 13, 1–43.

    Sellers, W.F. (1930) The identity of Zenillia blanda O.S. and Zenillia virilis A. & W. with notes on Zenillia blandita Coq. (Diptera, Tachinidae). Annals of the Entomological Society of America, 23, 569–576.

    Sellers, W.F. (1943) The Nearctic species of parasitic flies belonging to Zenillia and allied genera. Proceedings of the United States National Museum, 93 [= No. 3157], 1–108.

    Shima, H. (1996) A systematic study of the tribe Winthemiini from Japan (Diptera, Tachinidae). Beiträge zur Entomologie, 46, 169–235.

    Smith, H.E. (1912) A contribution on North American dipterology. Proceedings of the Entomological Society of Washington, 14, 118–127.

    Smith, S.M., Frankenhuyzen, K. van, Nealis, V.G. & Bourchier, R.S. (2002) Choristoneura fumiferana (Clemens), eastern spruce budworm (Tortricidae). In: Mason, P.G. & Huber, J.T. (Eds.), Biological control programmes in Canada, 1981–2000. CABI Publishing, Wallingford and New York, pp. 58–68.

    Tilles, D.A. & Woodley, N.E. (1984) Spruce budworm parasites in Maine: a reference manual for collection and identification of common species. United States Department of Agriculture, Forest Service, Cooperative State Research Service, Agriculture Handbook, 616, 35 pp.

    Thompson, W.R. (1953) The tachinid parasites of Archips cerasivorana Fitch. (2) Eusisyropa blanda O.S. (Diptera). Canadian Entomologist, 85, 393–404.

    Torgersen, T.R. (1985) Parasitoids and western budworm dynamics. In Sanders, C.J., Stark, R.W., Mullins, E.J. & Murphy, J. (Eds.), Recent advances in spruce budworms research. Proceedings of the CANUSA spruce budworms research symposium. Bangor, Maine, September 16–20, 1984. Canadian Forestry Service, Ottawa, pp. 104–105.

    Torgersen, T.R., Campbell, R.W., Srivastava, N. & Beckwith, R.C. (1984) Role of parasites in the population dynamics of the western spruce budworm (Lepidoptera: Tortricidae) in the Northwest. Environmental Entomology, 13, 568–573.

    Tothill, J.D. (1912) Systematic notes on North American Tachinidae. Canadian Entomologist, 44, 1–5.

    Tothill, J.D. (1913) Tachinidae and some Canadian hosts. Canadian Entomologist, 45, 69–75.

    Tothill, J.D. (1923) Notes on the outbreaks of spruce budworm, forest tent caterpillar and larch sawfly in New Brunswick. Proceedings of the Acadian Entomological Society [for] 1922, 8, 172–182.

    Townsend, C.H.T. (1908) The taxonomy of the muscoidean flies, including descriptions of new genera and species. Smithsonian Miscellaneous Collections, 51 (2 [= pub. 1803]), 1–138.

    Townsend, C.H.T. (1926) New holarctic Muscoidea (Diptera). Insecutor Inscitiae Menstruus, 14, 24–41.

    Vakenti, J.M., Cossentine, J.E., Cooper, B.E., Sharkey, M.J., Yoshimoto, C.M. & Jensen, L.B.M. (2001) Host-plant range and parasitoids of obliquebanded and three-lined leafrollers (Lepidoptera: Tortricidae). Canadian Entomologist, 133, 139–146.

    Walker, F. (1849) List of the specimens of dipterous insects in the collection of the British Museum. London, Vol. 4, pp. 689–1172.

    Webber, R.T. & Schaffner, J.V., Jr. (1926) Host relations of Compsilura concinnata Meigen, an important tachinid parasite of the gipsy moth and the brown-tail moth. United States Department of Agriculture, Department Bulletin, 1363, 1–31.

    Wilkes, A. (1946) The introduction of insect parasites of the spruce budworm, Archips fumiferana Clem., into eastern Canada. Canadian Entomologist, 78, 82–86.

    Wilkes, A. (1949) Notes on the transfer of budworm parasites from British Columbia to eastern Canada. Report of the Quebec Society for the Protection of Plants [for] 1945–1947, 30, 111–112.

    Wilkes, A. & Anderson, M. (1947) Notes on recovery of the introduced spruce budworm parasite, Phytodietus fumiferanae Rohw. in eastern Canada. Annual Report of the Entomological Society of Ontario [for] 1946, 77, 40–44.

    Wilkes, A., Coppel, H.C. & Mathers, W.G. (1949) Notes on the insect parasites of the spruce budworm Choristoneura fumiferana (Clem.) in British Columbia. Canadian Entomologist, 80 (1948), 138–155.

    Wilkinson, T.K., Landis, D.A. & Gut, L.J. (2004) Parasitism of obliquebanded leafroller (Lepidoptera: Tortricidae) in commercially managed Michigan apple orchards. Journal of Economic Entomology, 97, 1524–1530.

    Williston, S.W. (1889) The dipterous parasites of North American butterflies. In: Scudder, S.H., The butterflies of the eastern United States and Canada with special reference to New England. Published by the author, Cambridge. Vol. 3, pp. 1912–1924 + pl. 89.

    Winn, A.F. & Beaulieu, G. (1915) A preliminary list of the insects of the province of Quebec. Part II — Diptera (two-winged flies). Supplement to Report of the Quebec Society for the Protection of Plants, 7, 108–159.

    Wishart, G. (1945) Aplomya caesar (Aldrich), a tachinid parasite of the European corn borer. Canadian Entomologist, 77, 157–167.

    Wood, D.M. (1987) Tachinidae. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Volume 2. Agriculture Canada Monograph, 28, 1193–1269.

    Zeller, S.M. & Schuh, J. (1944) Diseases and insect pests of cane fruits in Oregon. Oregon Agricultural Experiment Station Bulletin, 418, 1–58.

    Zwolfer, H. (1961) A comparative analysis of the parasite complexes of the European fir budworm, Choristoneura murinana (Hub.) and the North American spruce budworm, C. fumiferana (Clem.). Commonwealth Institute of Biological Control, Technical Bulletin, 1, 162 pp.