This study’s main objective was to determine how the incidence of blister rust has changed since the mid-1900’s. Five separate regions of New Hampshire were surveyed in 1998 for the presence of white pine blister rust. Survey regions consisted of the north country, upper valley, southwest, southeast, and east central zones of the state. In each of these regions, approximately 20 sites were established and at each site, five subplots of 10 white pine trees were systematically sampled. To get comparable infection level data from past decades, we searched for statewide surveys done by foresters or researchers.
Acknowledgments
Examination and use of many manuals, particularly those done by the Division of Forests and
Lands and the USDA Forest Service in the early days of blister rust control, helped us develop this
report. Photographs were taken by Jennifer Bofinger, and collected from archive files at the Forest
Insect and Disease Section. Margaret Miller-Weeks (Forest Pathologist, USDA Forest Service), and
Mary Torsello (Forest Pathologist, USDA Forest Service) provided valuable technical advice and
editing skills. Special thanks go to Karen Bennett and the staff at UNH Cooperative Extension for
their time and attention to detail in publishing this report.
Finally, we would like to thank all those foresters and technicians from the early days who
quickly realized the threat blister rust posed to New Hampshire's early successional forest. Without
their knowledge and dedication, the Forest Insect and Disease Section would not be in a
position today to work on current forest health issues in New Hampshire.
About the New Hampshire Division of Forests and Lands
The Division of Forests and Lands is a part of the Department of Resources and Economic Development.
The Division's mission is to protect and promote the values provided by trees and
forests. To accomplish this goal the division manages state forest resources, provides information
and education, and provides leadership in the protection of New Hampshire forests.
Within the Division of Forests and Lands, the Forest Protection Bureau is responsible for providing
statewide leadership in forest protection. To meet this goal the bureau enforces forest harvesting
laws, provides state-wide monitoring and investigation of abiotic and biotic forest damaging
agents, and provides training and assistance to control wildland fires.
Within the Forest Protection Bureau, The Forest Insect and Disease Section is responsible for
providing current data on populations, trends and effects of forest insects, diseases, weather
events, or abiotic stressors. The insect and disease section also provides investigation, technical
assistance, and training with regard to current forest health indicators.
Introduction
At the turn of the 20th century, a little known fungal
disease called white pine blister rust, Cronartium
ribicola, (WPBR) spread from Russia westward into
Europe. At the same time in the eastern United
States, large quantities of white pine seedling were
needed for massive reforestation efforts.
New England foresters turned to forest nurseries in
France and Germany to fill this demand. Because
they didn't know about WPBR, some shipments into
New York were infected with what would become
the most severe disease of white pine in North
America. In 1909 the first outbreak was found in
Geneva, New York. By 1922, WPBR surveys in New
Hampshire showed areas with as much as 50% of
the pine stems infected (USDA, 1953).
popular means of ribes eradication.
It was thought that since this disease requires an alternate host
of Ribes spp. (gooseberries and currants), eradication of the
alternate host would prevent further spread of the deadly
WPBR. In 1917, New Hampshire prohibited the sale or planting
of any ribes species and started many aggressive ribes
eradication programs. By 1953 an estimated 7 billion board feet
of lumber production was lost due to this disease (USDA, 1953).
If those trees had survived, they would be mature trees today.
Based on current stumpage prices, the value to the landowner
would be $1.05 billion. The value to the state as a whole
through value added manufacturing and recreation would be
ten times that amount. Ribes eradication programs dominated
the forest pest arena from 1917 until the late 1970's. Ribes
plants were destroyed throughout the range of eastern white
pine with the use of herbicides and plant pulling techniques.
ground, hanging them in a tree dries
and kills the roots.
Starting in the late 1970's, serious questions
arose about the effectiveness of eradication
programs and enforcement of the ribes quarantine.
At the same time the white pine resource
in New Hampshire was maturing and the
prime ribes sites overtopped with hardwood
forests. Today there are no eradication programs,
the quarantine goes largely unenforced
and until this survey, we had little information
on the current status of blister rust infection
statewide.
method.
Disease Biology
The white pine blister rust disease (WPBR)
is a classic example of a tree rust fungi.
"Blister" escribes the effect on pine bark
after infection. "Rust" describes the physical
characteristics of the fruiting bodies on
host tree bark or leaves. In the case of
WPBR, the damage causing fungus is
Cronartium ribicola.
Like most other rust diseases, WPBR requires
two completely different host species
to complete its life cycle. In this case the
two hosts are eastern white pine and almost
any of the native ribes species (approximately
70 in North America).
The cycle begins with disease spores which attach themselves to needles of eastern white pine in
the late summer and early fall. These spores penetrate through stomata on the pine needles and
then start colonizing and expanding down into the twig. Within one to four years, the twig is
infected and the disease breaks out through the thin bark causing blisters and bark cracking.
Once the cambium is broken and killed, the tree response is usually pitching from the damaged
area. Once the disease has broken out through the bark, it travels at about two inches per year
toward the main stem of the tree. At this same time, a different type of spore is produced on the
surface of the white pine branch and is wind-blown in search of the alternate host. If the spores
find a ribes plant they attach and infect by spring and early summer. A series of spore development
stages take place on the surface of the ribes leaf and by late summer, there are spores capable
of infecting white pine. None of the different spore stages generated on the pine tree are
capable of infecting other pine trees. This isn't true of spores found on ribes plants. These spores
do travel to other ribes and colonize.
(ADD DIAGRAMS)
Field Identification
The first sign a pine is infected with WPBR is yellowing
of a group of needles. This yellowing will
spread to all of the needles on the branch. This first
stage is called "lagging". Once the disease reaches
the main branch, WPBR manifests itself in much the
way poison ivy does on humans. Pockets of yellow,
slimy spores grow under the thin bark until the bark
ruptures and the spores ooze out. After a period of
several years the disease finally reaches the main
stem of the tree.
At this point, the cambium is killed around the
branch collar and cambium mortality progresses
around the tree until the tree completely dies. The
larger and more vigorous trees tend to resist the disease for many years. This creates a canker, a
flattened dead area along the bole, with a dead branch stub in the middle. Moderate pitching is
usually associated with the outer edge of the canker.
branches which turn yellow. This symptom is
called "flagging"
Wildlife such as porcupines are attracted to this disease-sweetened area of the bark and cause
additional wounding. Blister rust can be mis-identified in the field because heavy pitching has
long been associated with this disease. Blister rust cankers, in many cases, pitch very little in
comparison with mechanical wounds or pine canker (Caliciopsis pinea) infections. If the pitching is
internodal, thus not associated with a branch, it isn't a sign of blister rust. To make a positive
field identification, it is important to find evidence of the branch that the disease infected.
sapling with a severe basal infection.
Note the branch the disease entered on
and the infectious spores in a target
shape on the stem.
been chewed by a porcupine. Many
small animals find the newly infected
bark irresistible.
necessary to climb this white pine tree
to point out the blister rust infection.
Survey Methods
Five separate regions of New Hampshire were surveyed in 1998 for the presence of white pine
blister rust. Each separate survey region encompasses eight to 12 towns for a total of 50 towns
surveyed. Survey regions consisted of the north country, upper valley, southwest, southeast, and
east central zones of the state. In each of these regions, approximately 20 sites were established
and at each site, five subplots of 10 white pine trees were systematically sampled.
(ADD MAP DIAGRAM)
This method provided approximately 1000 white pine trees in each zone for analysis. Approximately
equal amounts of seedling/saplings, pole timber, small sawlog, and large sawlog size trees were
sampled within each zone. Sites were located randomly by driving roads transecting through the
zone. To qualify as an appropriate survey site, the forest had to be more than 90% stocked with
eastern white pine, larger than two acres, and show no signs of timber harvesting since the stand was
established.
Results
A total of 5195 white pine stems were examined for WPBR. Trees were located on 104 different sites
grouped in 5 different regions of New Hampshire. One hundred twenty three (2.4%) of the 5195 trees
examined were identified as having at least one bole or branch canker of white pine blister rust.
The frequency of WPBR in each region was determined by calculating the percent of all sites
visited with at least one infection. Figure 1 shows that the north region had the highest percentage
of plots infected with WPBR. The southwest region of New Hampshire had the lowest percentage
of diseased plots.
(ADD GRAPH DIAGRAM)
The north region of New Hampshire, extending from Littleton to Lancaster, had the highest
incidence of infection (Table 1). 7.2% of the 1000 trees inspected in that zone had at least one
WPBR canker. The southwest region of New Hampshire had the smallest percentage of infection.
Just 0.3% of the surveyed stems were infected. The size class with the highest incidence of infection
was the large sawlogs. Trees over 14 inches at diameter at breast height (dbh) had a statewide
infection incidence of 3.2%. However, there is no significant difference between any of the
classes.
Table 1-Incidence of Infection: Percent Infection by Size Class for Each Region Surveyed
Region: North
Seed/Sapling: 6.2
Pole size: 6.6
Small sawtimber: 10.2
Larger sawtimber: 9.4
All size classes: 7.2
Region: Upper Valley
Seed/Sapling: 2.7
Pole size: 2.6
Small sawtimber: 2.4
Larger sawtimber: 3.6
All size classes: 2.7
Region: Southwest
Seed/Sapling: 0.8
Pole size: 0.4
Small sawtimber: 0.0
Larger sawtimber: 0.0
All size classes: 0.3
Region: Southeast
Seed/Sapling: 2.3
Pole size: 0.6
Small sawtimber: 0.8
Larger sawtimber: 1.8
All size classes: 1.2
Region: Central
Seed/Sapling: 0.3
Pole size: 0.9
Small sawtimber: 0.5
Larger sawtimber: 0.4
All size classes: 0.6
Total Region: Total of North, Upper Valley, Southwest, Southeast, Central
Total Seed/Sapling: 2.2%
Total Pole size: 2.4%
Total Small sawtimber: 2.4%
Total Larger sawtimber: 3.2%
Total All size classes: 2.4%
Discussion
This study's main objective was to determine how the incidence of blister rust has changed since
the mid-1900's. To get comparable infection level data from past decades, we searched for statewide
surveys done by foresters or researchers. Surprisingly, we found no previous statewide
inventory data on the amount of infection. However, we did find anecdotal information from old
newspaper articles and local survey reports such as "The Hurlin Plantation, A Case Study," (S.H.
Boomer, L.E. Newman 1950) and regional reports like White Pine Blister Rust Control, The Annual
Report (USDA Bureau of Entomology and Plant Quarantine, 1953).
Within these reports, the incidence of infection on live trees ranged from 20% to 80%. Based on
our survey results and the data from early to mid 20th century reports we have far less blister rust
incidence of infection than we had in the past. However, the results of this survey suggest the
disease is still well distributed in the northern regions of the state, yet less severe within each
timber stand. Live blister rust cankers in the north country occurred on 85% of all sites, while in
the southwest they occurred on just 16.7% of the surveyed sites.
These results are consistent with a 1960 study done by John Charlton of the USDA Forest Service
(Charlton 1963). Charlton published the results of a study on the different climatic zones of the
northeast and the probability of infection from WPBR in each zone. He divided the northeast
United States into high, moderate and low hazard zones based on mean nocturnal summer
temperatures and regional air saturation levels.
WPBR spore production and longevity depends on cool temperatures and high amounts of
moisture, so the cooler and moister regions of New Hampshire received the highest hazard rating.
When comparing our results by region to these hazard zones, we discovered the north region of
the state with the coolest nocturnal temperatures and highest amount of precipitation had the
highest amount of infected white pine trees. The second highest amount of blister rust infection
occurred in the upper valley region and this area is also located in the high hazard zone. It is
interesting that our two least infected regions both occurred in the moderate hazard zone (southwest
and central NH), while the moderately infected region fell in the low hazard rating zone
(southeast).
While blister rust infection is still common in many regions of New Hampshire, the severity of the
incidence of infection has declined dramatically. The reason for this decline in incidence of infection
is due to a number of contributing factors. One factor is described by statewide forest inventory
data (Cullen and Leak 1988).
Cullen and Leak report 78% of the white pine resource in New Hampshire is in the sawlog size
class. In 1973 it was 57% and in the years prior to 1973 the percentage of large pine was even
smaller. This data is significant when combined with the fact that white pine trees with live
branches within 10 feet of the ground are most susceptible to infection. These facts combined
would suggest the majority of pine has grown out of the susceptible age class, so the infection rate
has dropped.
This mature forest condition can't be the only factor contributing to reduced infection statewide.
If it were, high amounts of blister rust among the small size classes would be evident. We believe
an important factor contributing to the low incidence of infection in small size classes is the
reduced number of ribes bushes throughout New Hampshire. It's theorized that if an alternate
host is less abundant then the chance of infection will be reduced.
Cullen and Leak, in 1988, describe New Hampshire forests as "...highly skewed towards fully and
overstocked stands. Herbaceous ground cover conditions and shrub layers, many times available
under poorly stocked and nonstocked stands, are unlikely to be available in any noticeable way."
It appears that the conversion of old agriculture land to mature hardwood stands is almost complete
in New Hampshire and the amount of preferred habitat for the alternate host has been
severely reduced. To support this further, an analysis of the preliminary data from the 1998
statewide forest inventory (done by the USDA Forest Service in 1996 and 1997) shows that just 10
plots of the 1000 statewide had ribes plants on them.
Six of the 10 infested plots were in the northern two counties where the highest incidence of
WPBR occurred. The other four were distributed equally throughout the larger southern portion
of New Hampshire. It's worth noting that five of the six ribes infested plots found in the north
had fir, spruce, elm, or black ash in the overstory. These species are generally associated with wet
sites. This data further supports what we already know about habitat requirements for ribes.
A third and related factor that could be contributing to reduced blister rust infections is the effectiveness
of ribes eradication programs. Ribes pulling and chemical spraying started in the 1920's.
and continued until the early 1970's. As many as 3000 acres per year in New Hampshire were
"ribes controlled". This large effort over such a long period of time surely reduced the numbers
of ribes plants in the population.
Results of this survey suggest there is no significant difference in infection incidence between size
classes. To better understand this, it is important to note that incidence of infection is equal to the overall amount of live infection at any one time in a forest. You must add new infections and subtract mortality to achieve incidence of infection. White pine blister rust invades through live branches close to the ground, so you would expect the smaller size classes to have higher rates of infection and higher rates of mortality, but not necessarily higher incidence of infection. If the mortality rate is close to infection rates then the incidence of infection is low.
On the sites we visited, we believe the mortality rates, though very low, are offsetting the infection
rate in the smaller size classes. Conversely the large, thick-barked, well-established pines
probably survive with the canker much longer than suppressed, densely stocked sapling stands,
giving us higher than expected incidence of infection figures for those large trees.>
Outlook For the Future
It's fair to suggest that if prime habitats for ribes continues to decline, no new ribes are introduced,
and most of New Hampshire's pine resource continues to grow out of the most susceptible
size classes, the statewide incidence of white blister rust will remain at current low levels. However,
there will be rare occasions in the north where pockets of pine regeneration receive heavy
amounts of infection.
It's generally recommended that foresters in the high hazard zones regenerate white pine under
an existing overstory. The overstory helps prevent moisture build-up on the needles which is
vital to the formation of blister rust. The Minnesota Department of Natural Resources suggests
the worst practice is to manage young pine in small openings at the base of slopes or in low areas
for the very same reason. If the forester, in high hazard zones, can afford to enter the stand shortly
after the final overstory removal of the previous stand to thin out any infected poles, and prune as
many as 200 of the best stems, the stand should be maintained as pine.
If low stocking levels, poor site index, or an exceptional infection rate persists after thinning, the
stand may need to be converted to a more appropriate forest type. In general, with proper white
pine silviculture and an understanding that some mortality in the dense seedling/sapling stands
contributes to improving the health of residual growing stock, white pine blister rust should not
be a limiting factor for foresters interested in pine management throughout New Hampshire.
Literature Cited
Boomer S.H. and L.E. Newman 1950 The Hurlin Plantation, A Case Study. Bureau of Entomology
and Plant Quarantine, Division of Plant Disease Control. Concord, NH.
Charlton, John W 1963 Relating Climate to Eastern White Pine Blister Rust Infection Hazard.
Eastern Region, Forest Service, USDA, Upper Darby, PA.
Cullen, J.B. and William Leak 1988 New Hampshire's Timber Resource: Past-Present-Future.
NH Division of Forests and Lands, DRED.
Leak, William B., J.B. Cullen and Thomas S. Frieswyk June 1995 Dynamics of White Pine in New
England. USDA, Forest Service, Northeastern Forest Experiment Station. Research paper NE-699.
USDA, Agriculture Research Service, Bureau of Entomology and Plant Quarantine 1953 White
Pine Blister Rust Control Northeastern Region. Greenfield, MA.
