Invasive Species and the Problems They Pose:

The American Chestnut

Buchanan, Sarah C.
April 17, 2015

Abstract
Species are continuously being transported to new locations and
becoming invasive species. While invasive species are not always a
bad thing, they can have detrimental outcomes. Castanea dentata, the
American chestnut, is one example of how invasive species can hurt a
population. The fungus Cryphonectria parasitica, or the chestnut blight
wiped out around 4 billion trees after arriving in the United States from
its native home of Asia. Much is being done to save the American
chestnut and reintroduce it back into its natural habitat but there are

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many other challenges it still faces. Despite the challenges, though,
the future looks bright.
From the moment man set out to explore new lands he had the
desire to collect new exotic plants and animals as a way to
demonstrate his explorations. By collecting these new species, though,
explorers removed them from areas they were isolated in for millions of
years. Once brought to the explorers home, the species is considered
invasive and in order to survive the invasive species may alter the
evolutionary pathway of native species. Competitive exclusion along
with, niche displacement, hybridization, introgression, predation, and
possibly extinction may occur. People living today see the results of
invasive species, both the good and the bad (Mooney and Cleland,
2001).

Today species are continuously being transported to new

locations and of the large number being moved only a fraction ever get
established in the new area. Of the species that do establish
themselves, only one percent causes a problem. In many cases the
invasive species just adds to the areas species richness. In New
Zealand, for example, there are now as many invasive plant species as
there are native species. Another example is Arizona and Montana.

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They at one time had no fish species in common and now they share
33 of the same species (Mooney and Cleland, 2001).

Invasive species, when they do establish themselves in a new

environment, are able to adapt very quickly. This is not always a good
thing. An invasive species could possibly out compete or become the
predator of the native species. The native species can be completely
eliminated from the area if the invasive species causes a threat, or in
some cases, be driven to extinction. Invasive species have become the
second overall risk to endangered species in the United States
(Mooney and Cleland, 2001) One such species put at risk by an
invasive species is Castanea dentata or the American Chestnut (Elliott
and Swank, 2008).

The American chestnut tree was once a widespread species in

the eastern forests of the United States and was once the most
abundant tree in the forests of the Appalachian Mountains (Adams and
Lipford, 1992). At one point the American chestnut had a natural range
of 800,000 km2 and covered 50% of the land if stood on (Jacobs,
2007), ranging from Main to Alabama and westward toward Mississippi
(Barakat et al., 2009). It was an important resource for the American

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people and gave a unique ecological niche to the forests it inhabited
(Jacobs, 2007). It grew rapidly and provided food and habitat for
wildlife and for people it provided rot-resistant lumber, tannins to
process leather, fuel, firewood, and nuts. Considering the American
chestnuts many uses and quick growing time many referred to it as
the perfect tree (Barakat et al., 2009).

The American chestnut was the perfect tree until the pathogen
Cryphonectria parasitica, commonly known as Barr or the chestnut
blight, wiped out nearly all of them in the early twentieth century
(Jacobs and Severeid, 2004). In 1904 the chestnut blight was first seen
in the Bronx Zoological Park in New York City. The pathogen was
introduced to the United States by Japanese or Chinese chestnut trees
brought over from Asia to be nursery stock plants (Barakat et al.,
2009). By the 1950s the blight had spread to nearly all the American
chestnut trees and by the 1960s had killed around 4 billion trees.
Understandably the trees now are considered endangered in their
natural range, Canada, and in several US states (Jacobs, 2007)

The pathogen kills the tree by infecting stem tissues and choking
the portion of the tree above ground (Barakat et al., 2009). Symptoms
common with the chestnut blight are bark cankers, wilting of foliage,

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and the growth of epicormic sprouts straight below the canker. The
cankers are not easily seen on mature trees and are most easily seen
on young trees when the bark is wet (Anagnostakis, 1987). After the
tree has died the roots of the tree can continue to survive. For years
the tree will send up sprouts, which will eventually get infected, too,
and the process will start again. Today, the once great trees resemble
shrubs, because of the multiple shoot coming out, more so than the
giant trees we once knew them as (Barakat et al., 2009).

The chestnut blight, though devastating, was not the first threat
to the American chestnut tree. In the early 19th century the fungus
Phytophthora cinnamomi, otherwise known as ink disease, arrived in
the United States from Asia and caused major damage to the southern
portion of American chestnut trees (Clark et al., 2014). Ink disease
killed the American chestnut trees, and hundreds of others (Rhoades et
al., 2003), by rotting the roots of the plants (Crandall et al., 1945). The
fungus first travels through the soil as spores and creates lesions on
the roots before rotting (Jacobs, 2007). Root rot caused by ink disease
may now be creating a limitation to the creation of blight resistant
hybrid American chestnuts. It was found that ink disease is more
prevalent in wet, slightly and very compact soils. Knowing this it is

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important to consider placement of American chestnut trees. When
planting trees that are resistant to the chestnut blight it is best to plant
them in soils that do not promote root rot by ink disease (Rhoades et
al., 2003). It has also been note that Chinese chestnut exhibits some
resistance to ink disease and it has been suggested that the selection
of the resistant gene in cross breeding would be beneficial (Jacobs,
2007).

Many nonnative insects also pose a problem for the American

chestnut. The oriental gall wasp, Dryocosmus kuriphilus, is native to
China and is notorious for laying its eggs in vegetative and floral buds
of chestnut trees. Eggs that have been laid can cause branch dieback,
fruiting delay, and potentially death to the chestnut. In the United
States the oriental gall wasp was first seen in the 1970s and it spread
very quickly to the American chestnuts (Jacobs, 2007).

Other insects that cause a hazard to the American chestnut are

the gypsy moths (Lymantria dispar) and Ambrosia beetles
(Xylosandrus saxeseni). Both of these insects were once natives of Asia
and were introduced to the United States only to cause problems.
Gypsy moths will most likely interfere with the reintroduction of the
American chestnut because it has lately damaged more hybrids than it

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did pure American chestnut trees. Ambrosia beetles, too, will be a
potential threat because they are known to hurt Chinese chestnut trees
(Jacobs, 2007).

The reestablishment

of the American chestnut has been given

great attention over the last century and has been a goal for many tree
breeders (Clark et al., 2014). The breeding program has been kindly
supported for many years by federal, state, private, and non-profit
organizations to initiate the largest forest restoration project. If it can
be done, the restoration of an almost pure American chestnut would
give back a unique ecological niche that was once abundant (Jacobs,
2007). When the chestnut blight was first seen in the United States,
the now nonoperational United States Department of Agriculture
Bureau of Plant Industry began trying to save the tree by crossing the
American chestnut with resistant Asian chestnuts. In the 1940s the
operations were then reassigned to the Connecticut Agricultural
Experiment Station. Unfortunately the work was largely unsuccessful.
It was not until 1986 that serious research started back up again when
the backcross breeding method was proposed (Clark et al., 2014).

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The procedure for backcross breeding requires a donor parent,
the individual with the gene of interest (in this case an Asian chestnut
with blight resistance), and a recurrent parent, the individual that can
be made better by adding the gene of interest (the American
chestnut). The American and Asian chestnut are crossed and the
offspring of the two are then crossed back with the American chestnut.
Any offspring of this cross that exhibit resistance to the chestnut blight
is then crossed back with the American chestnut. This process is
repeated as many times as necessary until the end result is an
American chestnut tree with chestnut blight resistance (Advanced
Backcross Breeding, 2015).

Currently there are three major organizations trying to fight the

chestnut blight and ink disease. The first and oldest of these
organizations is the Connecticut Agricultural Experiment Station
(CAES). The CAES primarily works on creating hybrids using
backcrossing between the American chestnuts and Japanese and
Chinese chestnuts to fight against the chestnut blight, ink disease, and
also the Asian gall wasp (Clark et al., 2014).

The second organization is the American Chestnut Cooperators

Foundation (ACCF). Founded in 1980 they mainly try to fight the

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chestnut blight by using hypovirulence for blight control with a small
amount of blight resistant crossing (Clark et al., 2014). Hypovirulence
means they are infecting the blight with a virus that will drastically
weaken it. The tree, though, is not hurt by the virus and is able to fight
back against the blight. The tree is able to block the fungus with layers
of hard tissue and builds a barrier around the infection. The barrier is
and ugly knotty mass but the tree survives and that is what is
important (Hypovirulence-Infecting the Infection, 2015).

Hypovirulence was first seen occurring naturally in nature in

Switzerland. European chestnuts were also suffering from chestnut
blight and Dr. Jean Grente noticed a tree with healing infections. He
isolated the fungus from this tree and applied it to other European
chestnut trees, which is why they are alive today. The only draw back
to this in the United Stats is that in order for the blight to be infected
the funguses must join together and they have to share vegetative
compatibility genes. If one gene is different fusion will probably not
take place. Hypovirulence worked very well in Europe because there
are fewer types of vegetative compatibility than in the United States
(Hypovirulence-Infecting the Infection, 2015).

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The third and most well know organization fighting the blight is
the American Chestnut Foundation (TACF). Founded in 1983 the main
facility of TACF is located in Meadowview, Virginia. This primary
orchard produces the most advanced trees for breeding whose
seedlings are hypothesized to have blight resistance most similar to
that of the Chinese parent tree and traits most morphologically similar
to that of the American parent tree. The foundation has also started
testing for ink disease resistance, as well (Clark et al., 2014).

Reintroduction of a blight-resistant American chestnut is

projected for the near future. In many cases reintroduction will mean
intermixing the hybrid chestnut with other hardwood species.
However, there is a question of how well the hybrid chestnut will do
being so near other trees compared to how well the pre-blight chestnut
did due to competition with the other hardwood trees. A study was
done where American chestnut was planted in the same area as black
walnut (Juhlans nigra) and northern red oak (Quercus rubra) in a region
of southwestern Wisconsin where there is no indication of chestnut
blight. The American chestnut displayed a greater height (77% more)
and a greater diameter at breast height (140% more) than both the
black walnut and the northern red oak. The American chestnut also

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displayed better stem formation than the other two trees. These
results confirm that reintroduced trees should do just as well at
growing as pre-blight trees did and they are expected to show similar
silvical characteristics to the trees that grew in the study (Jacobs and
Severeid, 2004). Intervention still might be needed, though, because
the rapid initial growth of the American chestnut may possibly
suppress other hardwood species. The American chestnut though, has
reportedly been outcompeted by other seedling species if grown
initially from seed in clear-cut areas. Burning programs or herbicide
applications may have to be implemented to control competition in
reintroducing the American chestnuts to clear areas (Jacobs, 2007) .

There are many hindrances threatening the reestablishment of

the American chestnut tree and chief among these is the acceptance
of the hybrid chestnut tree (Jacobs, 2007). The offspring of backcross
breeding the different chestnut trees cannot actually be called a pure
American chestnut, though. These hybrid offspring may even be
classified one day as a new species. This may cause some uncertainty
when trying to introduce the trees back into the American chestnuts
natural habitat because some people will raise questions concerning
the effects of a new species on an ecosystem. The hybrid trees should
cause no problem to the ecosystem, though, because they will be

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made up of more than 90% of American chestnut tree and
phenotypically no one will be able to see a difference between the
original American chestnut and the hybrid. When all is said and done
the hybrid trees will be the American chestnut, just with added
resistance to the chestnut blight, and there should be no problem of
someone saying a non-native species is being introduced (Jacobs,
2007).
Now we ask the question, why is so much effort going in to save
a species that faces so many challenges and requires so many
resources? A sense of nostalgia, maybe, for the once great giant, or
maybe for its iconic status it once had of being the perfect tree.
Whatever the reason its successful reestablishment will provide great
ecological value and if success will give helpful insight and hope in the
fighting of other problems caused by invasive species. It does take a
lot of time but with some patience the American chestnut tree might
have the capability of returning as a dominant canopy tree in the
Appalachian Mountains and other eastern parts of the United States
(Clark et al., 2014) and despite all of the challenges that go with trying
to reestablish the American chestnut, the knowledge that the largest
ecological conservation story of our time was a success makes it all
worth it (Jacobs, 2007).

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Works Cited
Adams, H. S. and Lipford, M. L. 1992. The Impact of Human Activities
on the Upland
Forests of Western Virginia. Virginia Journal of Science, 43:121131
"Advanced Backcross Breeding." Plant and Soil Sciences ELibrary. N.p.,
n.d. Web.
2015.<http://passel.unl.edu/pages/informationmodule.php?
idinformationmodule=959723462&topicorder=2&maxto=7>.
Anagnostakis, S. L. 1987. Chestnut Blight: The Classical Problem of an
Introduced
Pathogen. Mycologia, 79:23-37
Barakat, A., DiLoreto, D. S., Zhang, Y., Smith, C., Baier, K., Powell, W. A.,
Wheeler, N.,
Sederoff, R., and Carlson, J. E. 2009. Comparison of the
transciptomes of American chestnut (Castanea dentata) and
Chinese chestnut (Castanea mollissima) in response to the
chestnut blight infection. BMC Plant Biology, 9:51
Clark, S. L, Schlarbaum, S. E., Pinchot, C. C., Anagnostakis, S. L.,
Saunders, M. R.,
Gundy, M.T., Schaber, P., McKenna, J., Bard, J. F., Berrang, P. C.,

14
Casey, D. M., Casey, C. E., Crane, B., Jackson, B. D.,
Kockenderfer, J. K., Lewis, R. F., MacFarlane, R., Makowski, R.,
Miller, M. D., Rodrigue, J. A., Stelick, J., Thornton, C. D, and
Williamson, T. S. 2014. Reintroduction of American chestnut in
the national forest system. Journal of Forestry, 115:502-512
Crandall, B. S., Gravatt, G. F., and Ryan, M. M. 1945. Root disease of
Castanea species
and some coniferous and broadleaf nursery stocks, cuased by
Phystophthora cinnamomi. Phytopathology, 35:162-180
Elliott, K. J. and Swank, W. T. 2008. Long-term changes in forest
composition and
diversity following early logging (1919-1923) and the decline of
American chestnut (Castanea dentata). Plant Ecology, 197:155172
"Hypovirulence-Infecting the Infection." The American Chestnut
Foundation
Virginia Chapter. The American Chestnut Foundation, n.d. Web.
2015. <http://www.vatacf.org/Hypovirulence.shtml>.
Jacobs, D. F. 2007. Toward development of silvical strategies for forest
restoration of
American chestnut (Castanea dentata) using blight-resistant
hybrids. Biological Conservation, 137:497-506
Jacobs, D. F. and Severeid, L. R. 2004. Dominance of interplanted
American chestnut
(Castanea dentata) in southwestern Wisconsin, USA. Forest
Ecology and Management, 191:111-120
Mooney, H. A. and Cleland, E. E. 2001. The evolutionary impact of
invasive species.
PNAS, 98:5446-5451
Rhoades, C. C., Brosi, S. L., Dattilo, A. J., and Vincelli, P. 2003. Effect of
soil compaction
and moisture on incidence of Phytophthora root rot on American
chestnut (Castaea dentata) seedling. Forest Ecology and
Management, 184:47-54