Mycological Society of America
Identification of Armillaria Species from New Hampshire
Author(s): T. C. Harrington and D. M. Rizzo
Source: Mycologia, Vol. 85, No. 3 (May - Jun., 1993), pp. 365-368
Published by: Mycological Society of America
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Mycologia, 85(3), 1993, pp. 365-368.
? 1993, by The New York Botanical Garden, Bronx, NY 10458-5126
BRIEF
IDENTIFICATION
ARTICLE
OF ARMILLARIA
SPECIES
FROM
NEW
HAMPSHIRE
T. C. Harrington
Department of Plant Pathology, Iowa State University, Ames, Iowa 50011
AND
D. M. Rizzo
Department of Plant Pathology, University of Minnesota,
St. Paul, Minnesota 55108
In 1979 Anderson and Ullrich proposed that
Armillaria mellea sensu lato in North America
be recognized as 10 biological species (Anderson
and Ullrich, 1979). Although basidiome morphology was not compared, the biological species
could be identified by compatibility testing; single-basidiospore strains of a given biological spe?
cies generally changed their cultural morphology
when paired on artificial media (Korhonen, 1978).
Compatibility relationships and morphology of
these biological species have been further investigated in North America, Europe and elsewhere
(Watling et al., 1991). Six biological species found
in eastern North America have been related to
European species of Armillaria (Anderson et al.,
1980; Watling et al., 1991) or described as new
species (Berube and Dessureault, 1988, 1989).
Ar?
Because of difnculties in identification,
millaria species found in eastern North America
have not been extensively characterized, but indications are that they vary greatly in pathogenicity, in host specificity, and in the production
of rhizomorphs (Berube and Dessureault, 1988,
1989; Blodgett and Worrall, 1992a, b; Dumas,
1988; Gregory et al., 1991; Rizzo and Harring?
ton, 1993). We have identified six species of Ar?
millaria in New Hampshire. Herein we record
the hosts and distributions for five of these spe?
cies.
Basidiomes, mycelial fans, decayed wood and
rhizomorphs of Armillaria were collected wherever encountered in the state from 1983 to 1990.
These collections were not made systematically
but, rather, were haphazard collections, often as
part of other field studies (e.g., Rizzo and Har?
rington, 1988; Worrall and Harrington, 1988).
Most of the collections were made in the White
365
Mountain
National
Forest of central New
Hampshire (Grafton and Carroll Counties) and
the seacoast area of southeastern New Hamp?
shire (Rockingham and Strafford Counties).
Mycelial fans, basidiome tissue, or decayed
wood were axenically excised and placed on wa?
ter agar, 1% malt extract agar (MEA), or MEA
with 100 Mg/ml streptomycin sulfate and 4 fig/
ml benomyl
50WP added after autoclaving
(BSMA) (Harrington et al., 1992). Rhizomorphs
were surface-sterilized in 70% ethanol for a few
seconds and rinsed in sterile distilled water prior
to plating. Haploid strains derived from single
basidiospores were obtained by dilution plating
of spore masses from spore prints (Harrington et
al., 1992). Isolation plates were incubated at 2025 C and subcultures stored at 4 C on MEA.
All field isolates (putatively diploid) and single-basidiospore strains (putatively haploid) were
identified to species by pairing on MEA with
haploid tester strains of known Armillaria spe?
cies (Harrington et al., 1992; Rizzo and Har?
rington, 1992). The tester strains (Table I) were
selected for flufry morphology and consistently
changed morphology after pairing with diploid
isolates or compatible haploid strains of the same
species.
Five species of Armillaria were identified from
74 collections: A. calvescens Berube & Dessureault, A. gallica Marxmiiller & Romagn., A.
mellea (VahlrFr.) Kummer, A. ostoyae (Ro?
magn.) Herink, and A. sinapina Berube & Dessureault (Table II). The 74 collections were associated with 19 host species (five coniferous
species and 14 hardwood species). Two collec?
tions were not compatible with the testers and
were not identified.
366
Mycologia
Table I
Haploid tester strains used to identify diploid field isolates and haploid strains of Armillaria
a From J. J. Worrall.
b From J. B. Anderson.
Armillaria ostoyae was the most commonly
encountered species (Table II); it was collected
52 times, from the northern part of the state
south to the seacoast area, and from five coniferous and five hardwood species. Twelve ofthe
collections were from basidiomes (either singlebasidiospore isolations or isolations from basidiome tissue), and 40 were from mycelial fans,
decayed wood, or rhizomorphs. The majority of
the collections were from red spruce (Picea rubens) or balsam fir (Abies balsamea). In these
and other host species, the fungus usually appeared to be pathogenic because it occurred on
a living or recently killed tree. Occasionally, A.
ostoyae was found causing a root and butt rot of
living trees, i.e., there was decay of the central
xylem of roots and the butt without apparent
colonization of the cambial region (Rizzo and
Harrington, 1988).
All eight records of A. mellea were from basidi?
omes found in four mixed hardwood-conifer
stands in the seacoast area. No samples of A.
mellea were collected from inland sites. Four of
the basidiomes were collected from well-decayed
stumps or buried roots that could not be iden?
tified to species (Table II). In four other cases,
A. mellea was found killing roots and causing
dieback on living Pinus strobus, Acer sp., Betula
populifolia, or Quercus rubra. The apparent restricted range of A. mellea to the milder seacoast
area is consistent with other records of its occurrence in milder regions of Michigan (Proffer
et al., 1987), New York (Blodgett and Worrall,
1992a) and Europe (Kile et al., 1991).
Armillaria sinapina was found three times,
twice in central New Hampshire and once in
extreme northern New Hampshire (Coos County). Two isolations were from rhizomorphs growing through advanced hardwood decay caused
by other fungi. A third isolate was from a basidiome on a downed, well-decayed birch log.
Armillaria gallica was found nine times on
seven host species (Table II). It was recorded
from five sites in the seacoast area and two sites
in southcentral New Hampshire (Merrimack and
Grafton Counties). Records of A. gallica in New
York (Blodgett and Worrall, 1992a) were also
primarily from the central and southern portions
Brief
Article
367
Table II
Number of collections
and host of origin of Armillaria
of that state. Six ofthe isolates of A. gallica were
from basidiomes on or under a living tree, on
which it may have been pathogenic. Pathogenic
colonization was likely in three cases where iso?
lates were obtained from mycelial fans under the
bark of two chlorotic Malus sp. from apple orchards and from a dead, ornamental Elaeagnus
angustifolia.
Armillaria calvescens was collected twice from
two host species (Table II) in the northern hardwood forest type of the White Mountains. Both
isolates were from mycelial fans on living trees,
and the fungus appeared to be weakly pathogenic
(attacking previously stressed trees).
A sixth species, A. gemina Berube & Dessu?
reault, not identified among the 74 isolates re?
ported here, was found at an intensively studied
site in central New Hampshire, at which an additional 130 Armillaria isolates were obtained.
Detailed observations of host relationships of A.
gemina, A. calvescens, and A. ostoyae at this site
were reported previously (Rizzo and Harrington,
1993).
As in many other surveys for Armillaria, an
accurate portrayal of the distribution and hosts
in New Hampshire was limited because of sam?
rhi?
pling bias. The abundance of basidiomes,
zomorphs and mycelial fans of Armillaria species
varies greatly among the species and among sites.
species identified in New Hampshire
Our search for basidiomes was largely confined
to the White Mountains and the seacoast area.
Other isolations were biased to the more pathogenic species, particularly A. ostoyae, and to red
spruce and balsam fir because we preferentially
sampled dying trees of these two host species in
other studies. Given these limitations, it is somewhat surprising that we have identified all six
Armillaria species of the northeast United States
in the small state of New Hampshire.
We thank J. B. Anderson and J. J. Worrall for providing some of the tester strains used in this survey.
Also, D. Hobbins, M. L. Fairweather, K. J. Smereka
and P. J. Zambino assisted in the collection of basidi?
omes and other samples. We thank N. A. Anderson,
R. A. Blanchette and J. J. Worrall for their helpful
reviews of the manuscript. This material is based upon
work supported by the U.S. Department of Agriculture,
CSRS under Agreement No. 86-FSTY-9-0186. Journal
Paper No. J-14951 of the Iowa Agriculture and Home
Economics Experiment Station, Ames, Iowa Project
No. 0159.
Key Words: Armillaria, distribution, hosts, species
identification
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