The Severity of Budworm-Caused Growth Reductions in Balsam Fir/Spruce Stands Varies with the Hardwood Content of Surrounding Forest Landscapes
The Severity of Budworm-Caused Growth Reductions in Balsam Fir/
Spruce Stands Varies with the Hardwood Content of Surrounding
Forest Landscapes
Elizabeth M. Campbell, David A. MacLean, and Yves Bergeron
Keywords: budworm impacts, defoliation, dendroecology, insect outbreaks
P
ERIODIC SPRUCE BUDWORM (Choristoneura fumiferana Clem.) (Lepidoptera: Tortricidae) outbreaks
occur about every 35 years in the boreal and mixedwood forest regions of eastern North America (Royama
1984, Jardon et al. 2003). During outbreaks, repeated annual
defoliation by budworm larvae reduces tree growth and
often kills its hosts, particularly balsam fir (Abies balsamea
[L.] Mill.), over vast areas (MacLean 1980, Hardy et al.
1986, Bergeron et al. 1995). Although budworm outbreaks
are natural forest disturbances that play a critical role in
directing large-scale ecological processes (e.g., forest succession and nutrient cycling), they also reduce timber supplies and generate substantial socioeconomic losses. From
1977 to 1987, 17–22% of the annual timber harvest level
was lost to spruce budworm in Canada (Sterner and Davidson 1982, Power 1991, National Forestry Database 1995).
During the spruce budworm outbreaks of the 1970s,
1980s, and early 1990s, aerial application of insecticide was
used throughout much of eastern Canada to reduce budworm damage and economic losses. Broad-spectrum chem-
ical insecticides (e.g., fenitrothion), as well as the biological
insecticide Bacillus thuringiensis (B.t.), were applied. Although chemical insecticides are no longer considered an
environmentally acceptable option for controlling insect
outbreaks in Canada, B.t., and the insect growth regulator,
Mimic, can be used to reduce budworm populations during
outbreaks to protect foliage and prevent host deaths. Past
management efforts to reduce the impact of spruce budworm outbreaks in eastern Canada have relied heavily on
the application of insecticides, but given concern about the
potential negative ecological impacts of spraying B.t., for
example, dramatic reductions of Lepidoptera that could
significantly alter forest insect communities and affect a
range of other species and ecosystem processes (Crawford
and Jennings 1989, Miller 1990), additional approaches to
reducing the economic impacts of budworm outbreaks are
necessary.
The fact that damage caused by spruce budworm outbreaks varies with forest species composition has long been
recognized. On the basis of these observations, researchers
Elizabeth M. Campbell, Centre d’Étude de la Forêt and Département des Sciences Biologiques, C.P. 8888, Succ. Centre-ville, Université du Québec à
Montréal, Québec H3P 3P8, Canada. Current address: Research Branch, British Columbia Ministry of Forests and Range, PO Box 9519, Stn Prov Govt,
Victoria, BC V8W 1N1, Canada—Phone: (250) 387-6712; Fax: (250) 387-0046; David A. MacLean, Faculty of Forestry
and Environmental Management, University of New Brunswick, PO Box 44555, Fredericton, New Brunswick E3B 6C2, Canada. Yves Bergeron, Chaire
industrielle CRSNG-UQAT-UQAM en aménagement forestier durable, Université du Québec en Abitibi-Témiscamingue, C.P. 700, Rouyn-Noranda, Québec
J9X 5E4, Canada.
Acknowledgments: This project was funded by National Science and Engineering Research Council (NSERC) and Quebec–Canadian Provinces University
Cooperation Program grants and through scholarships obtained by E.C. from the Sustainable Forest Management Network and the NSERC/Université du
Québec à Montréal/Université du Québec en Abitibi-Témiscamingue Industrial Chair in Sustainable Forest Management. We thank Wayne MacKinnon,
Canadian Forest Service (Fredericton) for geographic information system work, assistance with core collections, and stand data from 12 of his research sites.
Guillaume Sauzède and Peter Goulet helped with fieldwork. Vince Zelazny (New Brunswick Department of Natural Resources) provided the ArcView map
layer of ecoregions onto which we overlaid our sample plots. We gratefully acknowledge two anonymous reviewers whose comments helped to significantly
improve this manuscript.
Manuscript received April 18, 2006, accepted September 21, 2007
Copyright © 2008 by the Society of American Foresters
Forest Science 54(2) 2008
195
Abstract: Tree-ring data collected from balsam fir (Abies balsamea [L.] Mill.) in 31 spruce (Picea spp.)/firdominated stands in northern New Brunswick were used to evaluate how radial growth reductions during the last
spruce budworm (Choristoneura fumiferana Clem.) outbreak (1972–1992) varied according to the hardwood
content of surrounding forest landscapes, insecticide spray history and elevation. Stepwise regression analysis
was used to build 12 models describing relationships between mean percent growth reduction (i.e., deviation
from mean tree growth averaged over the lifespan of all trees in a stand) and the explanatory variables for years
when ⬎80% of stands had growth reductions (1976 –1978, 1982–1984, 1990, 1991, and 1994) and for three
growth reduction periods (1972–1980, 1981–1985, and 1986 –1993). The hardwood content of surrounding
forest landscapes entered into all significant regression models (P ⬍ 0.05), explaining 12– 41% of the variance
in budworm-caused growth reductions. Differences in growth reductions among stands were greatest during the
initial years of the budworm outbreak (1972–1980), particularly in 1976 when mean reductions were ⬃40% in
stands with ⬍50% hardwood and content and ⬃20% in stands in which surrounding hardwood content was
⬎50%. Relationships between variation in growth reductions among stands and elevation or insecticide spray
history were weak and less consistent than with landscape hardwood content. Our findings suggest that
increasing the hardwood content of forest landscapes could help to reduce the impact of outbreaks on timber
supplies. FOR. SCI. 54(2):195–205.
196
Forest Science 54(2) 2008
of defoliation calculated from aerial surveys) (e.g., Hardy et
al. 1986). Studies such as this one and others that quantify
stand-level impacts on the basis of measurements of individual trees (e.g., MacKinnon and MacLean 2004) better
reflect local budworm population densities and thus provide
an improved basis for understanding relationships between
forest composition, outbreak impacts, and the mechanisms
that drive budworm population dynamics. This information
could have significant implications for the way in which
forests are managed to reduce future socioeconomic losses
caused by budworm outbreaks.
Methods
Study Area
Thirty-one stands were sampled in northern New Brunswick. Most stands occurred within the Northern Uplands
bioclimatic ecoregion, but a few stands occurred in the
Highlands and the Central Uplands ecoregions (Figure 1).
Elevations ranged from 177 to 549 m, with a mean of
335 m. The Northern Uplands is climatically intermediate to
the colder Highlands and the warmer, wetter Central Uplands (TECWG 2003). Mean annual, sea (...truncated)