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How do Exotic Pests Impact Forested
Ecosystems and Landscape Trees in North America?

Gaston Laflamme
and Thomas Hofacker
Invasions of exotic forest pests
into North America (NA) likely began with the arrival of settlers
from Europe. These settlers had a direct and enormous impact in the
East and South as forests were cleared for cities, towns, and
agricultural purposes. So much so, that by the late 1800s, the
eastern and southern forests of NA bore little resemblance to the
native forests that the European settlers first encountered. This is
the backdrop against which invasions of exotic forest insects and
pathogens began to be recognized as the sciences of entomology and
plant pathology emerged on the continent.
Then came gypsy moth (Lymantria
dispar) and chestnut blight (Chryphonectria
parasitica). In the early 1900s, a massive (and unsuccessful)
gypsy moth eradication and suppression program began. Efforts to
deal with these introductions continue today, with millions of
dollars spent annually. While the gypsy moth was intentionally
imported into North America, chestnut blight was unintentionally
introduced on chestnut seedlings brought to
the New York Botanical Garden from Asia (Schumann 1991). The
disease spread rapidly, invading 80% of the host range in 10 years. American
chestnuts had virtually no resistance to this fungus, and over the
next 50 years, an estimated four billion trees were killed. Today,
only stump sprouts are produced and they are also killed by the
pathogen. The entire species, which once comprised
one-quarter of the eastern U.S. forest, became functionally extinct;
that is, the species still exists, but not in sufficient numbers to
provide a useful ecological structure or function in a significant
manner.
The disease caused considerable
economic loss in wood and nut production. In many areas, oaks have
replaced the chestnut. These oak trees were later severely
defoliated by gypsy moth. Even a century later, the impact of these
exotics continues. To learn more about chestnut blight, click on to
the nicely illustrated article at http://www.apsnet.org/online/feature/chestnut/top.html.
Hot on the heels of gypsy moth and
chestnut blight came white pine blister rust. This rust
disease is caused by the rust pathogen Cronartium ribicola,
involves three organisms: the fungus, the host (white pine), and an
alternate host (Ribes spp., currants and gooseberries). The
two most important white pine species damaged by the disease are Pinus
strobus in eastern NA and P. monticola in western NA. At
the end of the 19th century and into the next, owners of tree
nurseries derived a good income from producing ornamental trees.
Blister rust was well known in Europe, yet white pine seedlings were
imported for resale. Probably many lots of diseased seedlings were
shipped from Germany and France (Maloy 1997). The disease was first
found in NA in 1906 when infected Ribes were located at
Geneva, New York.
The white pine population has
dropped dramatically in many areas. In Minnesota, for example, the
old growth white pine forest covered 3,500,000 acres in 1837, in
1990, this dropped to less than 67,000 acres. From 1917 to 1970,
white pine dropped by 60% in Quebec, an area where it is an
important species. (Lavallée 1974). In the western US, the western
white pine is surviving on 5% of its former domain, and the
ecosystem has change dramatically (Neuenschawander et al. 1999).
It is difficult and even impossible
to reintroduce white pine in many regions because of the pathogen.
The disease also reduces the production of small fruits on the
alternate host (Ribes spp.). Again, the economic loss is
difficult to evaluate, but the impact is very high. You can find
more on white pine blister rust at http://www.rms.nau.edu/rust/
and http://ppathw3.cals.cornell.edu/tree/WPBRust.html
These cases illustrate the enormous
impact of exotic insects and diseases. Although wood losses have to
be considered important, chestnut blight is probably one of the
worst ecological disasters ever recorded. Periodic gypsy moth
outbreaks keep this introduction in the public's eye, but the impact
of chestnut blight is largely forgotten because the number of people
who remember what these magnificent forests and trees once is
rapidly dwindling. These two examples illustrate an important
pathway used by diseases: seedlings. The pathways above and below
are underlined.
Less severe effects are more
common. For example, butternut canker has basically
eliminated butternut from the southern parts of its range, but still
exists in some northern areas where it may have some natural
resistance. It may have been introduced
through infected seeds (Innes 1998). Dutch elm disease
has wiped out American elm in our urban forests, but elms are still
present in significant numbers in many native forests. The disease
arrived on elm logs imported for veneer from
Europe in 1930. The fungus was also isolated from wood boxes made of
European elm and used for shipping goods (Boyce J.C. 1961; Stipes
and Campana 1981). Dogwood anthracnose has restricted
American dogwood populations to forest edges in many higher
elevation sites in the eastern United States, but it still exists in
forests at lower elevations. Balsam woolly adelgid has killed
most mature Fraser fir in native forests, but it is widely grown as
a holiday tree. Mattson, (1997), estimated that only about 5% of the
exotic insects that attack woody plants in NA have produced "
well recognized, severe ecological impacts on the trees and
ecosystems that they occupy." This 5% figure may be a result of
a lack studies as well as a lack of impact. In fact, we can show
that hemlock woolly adelgid has killed many streamside
eastern hemlocks, Port-Orford cedar root disease has killed
many streamside Port-Orford cedars, and beech bark disease/beech
scale has killed many American beech, but the effects of these
trees deaths is largely unknown. Scleroderris
canker,
caused by the European race, has affected nursery growers in
localized areas because of the probation on the movement of pine
stock. Several of the diseases, such as beech bark disease, Dutch
elm disease, and European larch canker continue to spread in
NA. For more information, see Liebhold et al. (1995).
Likewise, the more subtle effects
caused by exotic insects and pathogens are even more unclear. More
than 400 species of exotic insects alone are known to attack woody
plants in NA. We can speculate that these exotics add additional
stress to trees and forests that must already cope with unnatural
stresses such as global warming, acid deposition, stratospheric
ozone depletion, soil compaction, fire control, human development,
and a host of other factors. Manion (1991) has postulated that this
additional stress may result in a decline death spiral from which it
is difficult for affected hosts to recover. We can also speculate
that these effects will often play out over relatively long time
frames compared to human life spans. This, indeed, seems to be the
case, but studies to determine and document the direct and indirect
effects of exotic invasions over time are urgently needed in order
to make a better case for adequate funding to help stem the invasive
tide.
Plant quarantine and international
trade
Canada, Mexico, and the
United States have tried to develop a common NA approach to deal
with potential exotic pests by participating in the North American
Forestry Commission and the North American Plant Protection
Organization. These groups discuss the distribution, regulatory and
mitigation measures, and research on exotics of common concern.
Literature cited
Boyce, J.C. 1961. Forest Pathology.
Third edition McGraw-Hill, New York. 572pp.
Innes, L. 1998. Siroccocus
clavigignenti-juglandacearum on butternut and black walnut
fruit. p. 129-132 In Laflamme, G.; J.A. Bérubé and R.C. Hamelin
eds. Foliage, Shoot and Stem Diseases of Trees. Proceedings of the
IUFRO WP 7.02.02 Meeting, Québec City, May 25-31, 1997. Information
Report LAU-X-122. Canadian Forest Service, Laurentian Forestry
Centre.
Lavallée, A. 1974. Une
réévaluation de la situation concernant la rouille vésiculeuse du
pin blanc au Québec. Forestry Chronicle 50:1-5.
Liebhold, A.M.; W.L. MacDonald; D.
Bergdahl and V.C. Mastro. 1995. Invasion by exotic forest pests: a
threat to forest ecosystems. For. Sci. Monogr. 30:1-49.
Maloy, O.C. 1997. White pine
blister rust control in North America: A case study. Annual Rev.
Phytopathol. 35:87-109.
Manion, P.D. 1991. Tree disease
concepts. Prentice Hall Inc., Englewood Cliffs, NJ. 402 pp.
Mattson, W.J. 1997. Exotic Insects
in North American Forests: Ecological Systems Forever Altered. P.
187-193. Proceedings of the Conference on Exotic Pests of Eastern
Forests, Nashville, Tennessee, April 8-10, 1977.
Neuenschawander et al. 1999. White
pine in the American West: A vanishing species- Can we save it. USDA
Forest Service RMRS-GTR-35. 20pp.
Schumann, G.L. 1991. Plant
Diseases: Their Biology and Social Impact. APS Press, St. Paul,
Minnesota, USA. 397 pp.
Stipes, R.J. and R.J Campana, eds.
1981.Compendium of elm diseases. APS Press. 96pp.
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