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EUROPEAN SPRUCE SAWFLY, Gilpinia hercyniae (Hartig) -- Diprionidae [also known as Diprion hercyniae
(Hartig)] (Contacts) A spruce (Picea
spp.) feeding insect native to most of Europe, the European spruce sawfly was
first noted as an accidental introduction in Canada in 1922 (Dahlsten &
Mills 1999). By 1930 a severe outbreak
was causing concern in the Gaspe Peninsula and by 1936 the sawfly threatened
to devastate the spruce forests of eastern Canada by extending its range
across all eastern Provinces and adjacent United States and causing severe
damage over an area of more than 10,000 sq. miles (McGugan & Coppel
1962). One of the most extensive projects undertaken
in classical biological control was begun against European spruce sawfly in
1933. Gilpinia hercyniae
was not at first distinguished from G. polytomum
(Htg.) and the Farnham House Laboratory in England (now known as CIBC) was
engaged to make large scale parasitoid collections from the latter species in
Europe. Initial studies revealed that
apart from the egg parasitoids, all other parasitoids develop so as to
overwinter in the host cocoon. This
simplified parasitoid collections in Europe to those stages of
development. A team of about 30
persons collected >1/2 million cocoons of G. polytomum
in Europe for shipment to Canada during 1932-40. Additionally more >1/2 million eggs and 31 million cocoons
of other spruce and pine feeding sawflies were shipped to supplement the
numbers of the less host specific parasitoid species available for field
release (Morris et al. 1937, Finlayson & Finlayson 1958). There were 96 species of primary and
secondary parasitoids obtained from these cocoon collections at the
Belleville Laboratory in Canada and a multiple introduction program involving
two egg parasitoids and 25 larval and cocoon parasitoids was initiated in
1933-51. The importation of a wide
variety of parasitoids from diverse hosts permitted the inclusion of several
sawfly pests as additional targets for some of the releases (McGugan &
Coppel 1962). The addition of an elaborate controlled
environment quarantine building was made at Belleville in 1936 allowed the
mass rearing of several of the imported European parasitoids. Dahlbominus fuscipennis,
a gregarious ectoparasitoid of prepupae, readily attacked cocoons in the
laboratory and was selected along with several other species for a massive
program of mass rearing for release.
The mass-rearing peaked in 1940 when a total of 221.5 million D. fuscipennis was released and by the end of the program in
1951 a total of 890 million directly imported or laboratory reared
parasitoids had been liberated (McGugan & Coppel 1962). Only 5 species of parasitoids out of 27
released became established over more than several generations, although four
additional species were recovered during the years shortly after
release. Three of the five species, D. fuscipennis, Exenterus amictorius
(Panz.) and E.
confusus Kerr, were widely
established only during the outbreak and have since not been recovered from G. hercyniae.
Although E. amictorius had little impact,
the other two species achieved variable but appreciable levels of parasitism
and have been credited with the decline of the outbreak in at least some
areas. Two other parasitoids, Exenterus vellicatus Cush. and Drino bohemica Mesn., never became
important until the collapse of the outbreak but have replaced the three
species present during the outbreak to maintain host population at low,
non-damaging densities. The epizootic of European spruce sawfly began
to decline in 1939-40, which coincided in the southern part of the range with
the occurrence of a nuclear polyhedrosis virus, Borrelinavirus hercyniae. This virus is thought to have been
accidentally imported and released in Canada along with the parasitoid
(Dahlsten & Mills 1999). It
spread rapidly to produce virus epizootics throughout most of the outbreak
range and by 1943 host population densities had declined to very light
infestations. Unlike other diprionid
sawflies, G. hercyniae larvae are not
gregarious and the rapid spread and subsequent impact of the virus was
attributed to its virulence (Bird & Elgee 1957). More recent studies in the Great Britain,
where G. hercyniae was accidentally
introduced from the European continent in 1968, indicate that birds play an
important role in virus transmission (Entwistle 1976). The importance of D. bohemica,
E. vellicatus and the NPV virus in maintaining the spruce
sawfly at low population densities in Canada has been inadvertently
demonstrated through chemical spray treatments aimed against spruce
budworm. Both in the early 1960's and
again in the 1970's sawfly population levels increased immediately following
the cessation of a 2-3 year spray treatment, due to the detrimental effects
of the spray on natural enemies, but declined after several generations as a
result of increased parasitism and the reappearance of the virus (Neilson et
al. 1971, Magasi & Syme 1961, Dahlsten & Mills 1999). Dahlsten & Mills (1999) pointed to several
interesting features of this successful biological control program. First the success of the accidental
introduction of the virus provides to date the most outstanding example of
the use of a pathogen in classical biological control. Its ability to control the sawfly
population in the absence of parasitoids has been demonstrated (Bird &
Burk 1961, Entwistle 1976) and in Canada it has persisted in the forest
environment since the initial introduction despite the low host densities
(Magasi & Syme 1981). The
multiple introduction program of parasitoids resulted in the establishment of
the two more effective and specific species, despite the release of a wide
range of potential competitors.
However, the continuous and large scale release of poorly adapted
parasitoids, which were later recovered only from other sawfly hosts, was
successful in inducing significant levels of mortality prior to the
introduction of the virus. For greater detail on the hosts, natural
enemies and biological control please refer to the following (Webber 1932,
Morris & Cameron 1935, Morris 1937, 1942; Baird 1939, 1940; Cushman 1940,
Balch et al. 1941, Peirson 1941, Reeks 1937, 1941, 1952, 1953; Peirson &
Nash 1940, Wilkes 1942, Balch & Bird 1944, Daviault 1944, Dirks 1944,
Briand 1949, Schaffner & Middleton 1950, Balch 1960, Griffiths 1961,
Turnbull & Chant 1961, Dowden 1962, Graham & Jones 1962, Martineau
1963, Neilson & Morris 1964, McLeod & Desalliers 1966, Rose &
Sippell 1966). REFERENCES: [Additional references may be found at: MELVYL
Library ] Baird, A. B. 1939. Biological control of insect pests in
Canada with special reference to the European spruce sawfly Gilpina polytoma Htg. Ent.
Soc. Ontario, 70th Ann. Rept. (1939):
51-56. Baird, A. B. 1940. A review of the spruce sawfly parasite
situation. Pulp and Paper Mag.,
Canada. Ann. Meeting, Woodlands
Sect., Canad. Pulp Assoc. Proc., Jan 1939:
43-4. Balch, R. E. 1960. The approach to biological control in
forest entomology. Canad. Ent.
92: 297-310. Balch, R. E. & F. T. Bird.
1944. A disease of the
European spruce sawfly, Gilpinia
hercyniae (Htg.), and its
place in natural control. Sci. Agric.
(Ottawa) 25: 65-80. Balch, R. E., W. E. Reeks & S. G. Smith. 1941.
Separation of the spruce sawfly from Gilpinia polytoma
(Hts.) (Diprionidae: Hymenoptera) and evidence of its introduction. Canad. Ent. 73: 198-203. Bird, F. T. & J. M. Burk.
1961. Artificially
disseminated virus as a factor controlling the European spruce sawfly, Diprion herycinae (Htg.), in the absence of introduced
parasites. Canad. Ent. 92:
228-38. Bird, F. T. & D. E. Elgee.
1957. A virus disease and
introduced parasites as factors controlling the European spruce sawfly, Diprion hercyinae (Htg.) in central New Brunswick. Canad. Ent. 89: 371-78. Briand, L. J. 1949. Notes on the spruce sawfly (Gilpinia hercyniae Htg.) and parasitism in the Park Reserve,
Quebec, area. Quebec Soc. Protect.
Plants Ann., 30th Rept. (1945-47):
180-81. Cushman, R. A. 1940. A review of the parasitic wasps of the
ichneumonid genus Exenterus
Hartig. U. S. Dept. Agric. Misc.
Publ. 354. 14 p. Dahlsten, D. L. & N. J. Mills. 1999. Biological
Control of Forest Insects. In: Bellows, T. S. & T. W. Fisher (eds.), Handbook of Biological Control: Principles and Applications. Academic Press, San Diego, New York. 1046 p. Daviault, L. 1944. Nouvelles observations sur la mouche a
scie europeene de l'epinette dans les forets du nord du St-Laurent. Quebec Soc. Protect. Plants Ann. Rept.
(1936-43): 39-43. Dirks, C. O. 1944. Population studies of the European spruce
sawfly in Maine as affected by natural enemies. J. Econ. Ent. 37:
238-42. Dowden, P. B. 1962. Parasites and predators of forest insects
liberated in the United States through 1960.
U. S. Dept. Agric. Agric. Handb. 226.
70 p. Entwistle, P. F.
1976. The development of an
epizootic of a nuclear polyhedrosis virus disease in European spruce sawfly, Gilpinia hercyinae. Proc.
1st Intern. Colloq. Invert. Pathol., Kingston, Canada. p. 184-88. Finlayson, L. R. & T. Finlayson. 1958. Notes on
parasites of Diprionidae in
Europe and Japan and their establishment in Canada on Diprion hercyinae
(Htg.) (Hymenoptera: Diprionidae).
Canad. Ent. 90: 557-63. Graham, A. R. & P. A.
Jones. 1962. Parasites and predators released against
forest insects in Canada, 1910-58.
Appendix I. In a Review of the
Biological Control Attempts Against Insects and Weeds in Canada. Commonwealth Inst. Biol. Control.
Contrib., Tech. Commun. 2: 129-75. Griffiths, K. J.
1961. The life history of Aptesis basizina (Grav.) on Neodiprion
sertifer (Geoff.) in
southern Ontario. Canad. Ent.
93: 1005-10. Magasi, L. P. & P. D. Syme.
1981. Chapter 52. Gilpinia
hercyinae (Hartig), European
spruce sawfly (Hymenoptera: Diprionidae), p. 295-97. In: J. S. Kelleher & M. A. Hulme (eds.),
Biological Control Programmes Against Insects and Weeds in Canada
1969-1980. Commonwealth Agric.
Bureau, London, England. 410 p. Martineau, R. 1963. Natural factgors regulating populations of
the European spruce swafly, Diprion
hercyniae (Hts.) in southern
Quebec. Canad. Ent. 95: 317-26. McGugan, B. M. & H. C. Coppel. 1962. A review of the
biological control attempts against insects and weeds in Canada. II.
Biological control of forest insects, 1910-1958. Commonwealth Inst. Biol. Control Tech.
Comm. No. 2: 35-216. McLeod, J. H. & G. Desalliers. 1966. Notes on Exenterus amictorius Panzer, an introduced parasite not established
on the Swaine jack-pine sawfly, Neodiprion
swainei Midd. Canad. Dept. Forest., Bimo. Prog. Rept.
22: 1-4. Morris, K. R. S.
1937. The prepupal stage in
Ichneumonidae, illustrated by the life-history of Exenterus abruptorius
Thb. Bull. Ent. Res. 28: 525-34. Morris, K. R. S. & E. Cameron. 1935. The biology of Microplectron fuscipennis Zett., a parasite
of the pine sawfly (Diprion sertifer Geoff.). Bull. Ent. Res. 26: 407-18. Morris, K. R. S., E. Cameron & W. F. Jepson. 1937.
The insect parasites of the spruce sawfly (Diprion polytomum
Htg.) in Europe. Bull. Ent. Res.
28: 341-93. Morris, R. F. 1942. Preliminary notes on the natural control
of the European spruce sawfly by small mammals. Canad. Ent. 74:
197-202. Neilson, M. M. & R. F. Morris. 1964. The regulation of
European spruce sawfly numbers in the Maritime Provinces of Canada from 1937
to 1963. Canad. Ent. 96: 773-94. Neilson, M. M., R. Martineau & A. M. Rose. 1971.
Chapter 39. Diprion hercyniae (Hartig), European spruce sawfly (Hymenoptera:
Diprionidae), p. 136-43. In: Biological Control Programmes Against Insects and Weeds in
Canada, 1959-1968. Commonwealth Inst.
of Biol. Control, Tech. Comm. No. 4.
266 p. Peirson, H. B. 1941. Control work on European spruce sawfly in
1940. Maine Forest Commr. Bien. Rept.
23. 10 p. Peirson, H. B. & R. W. Nash.
1940. Control work on European
spruce sawfly in 1939. Maine Forest
Serv. Bull. 12. 19 p. Reeks, W. A. 1937. Notes on the biology of Microplectron fuscipennis Zett., as a cocoon
parasite of Diprion polytomum Hartig. Canad. Ent. 69: 185-87. Reeks, W. A. 1941. On the taxonomic status of Gilpinia polytoma (Htg.) and G.
hercyniae (Htg.)
(Hymenoptera: Diprionidae). Canad.
Ent. 73: 177-88. Reeks, W. A. 1952. Establishment of Exenterus spp. (Hymenoptera: Ichneumonidae), parasites of
the European spruce sawfly, near points of liberation. Canad. Ent. 84: 76-86. Reeks, W. A. 1953. The establishment of introduced parasites
of the European spruce sawfly (Diprion
hercyniae (Htg.)) in the
Maritime Provinces. Canad. J. Agric.
Sci. 33: 405-29. Rose, A. H. & W. L. Sippell.
1966. Distribution of cocoon
parasites of the European pine sawfly, Neodiprion
sertifer (Geoffroy) in
southern Ontario. Canad. Ent.
98: 40-5. Schaffner, J. V., Jr. & W. Middleton. 1950.
The sawflies and horntails. In: F. C. Craighead (ed.), Insect Enemies of Eastern Forests. U. S. Dept. Agric. Misc. Publ. 657: 522-91. Turnbull, A. L. & D. A. Chant. 1961. The practice and
theory of biological control of insects in Canada. Canad. J. Zool. 39:
697-753. Webber, R. T. 1932. Sturmia
inconspicua Meigen, a
tachinid parasite of the gypsy moth.
J. Agric. Res. 45: 193-208. Wilkes, A. 1942. Notes on the laboratory propagation of
three European species of Exenterus
(Ichneumonidae), parasitic on sawflies.
Ent. Soc.Ontario 72nd Ann. Rept. 1941: 41-4. |