Giant Sequoia Insect, Disease, and Ecosystem Interactions1

Douglas D. Piirto2

Abstract: Individual trees of giant sequoia (Sequoia gigantea [Lindl.] afflict and kill other trees." Similarly Hartesveldt (1962)
Decne.) have demonstrated a capacity to attain both a long life and very concurred that "Sequoia's longevity and great size have
large size. It is not uncommon to find old-growth giant sequoia trees in
their native range that are 1,500 years old and over 15 feet in diameter at been attributed by nearly all writers, popular and scientific,
breast height. The ability of individual giant sequoia trees to survive over to its few insect and fungus parasites and the remarkable
such long periods of time has often been attributed to the species high resistance of the older trees to damage or death by fire.
resistance to disease, insect, and fire damage. Such a statement, however, is There is no record of an individual sequoia living in its
a gross oversimplification, given broader ecosystem and temporal interac-
tions. For example, why isn't there a greater representation of young-growth natural range as having been killed by either fungus or insect
giant sequoia trees throughout the mixed-conifer belt of the Sierra Nevadas? attack." Even as recently as 1991 Harlow and others (1991)
What other factors, in addition to physical site characteristics, limit giant stated: "Insects and fungi cause but minor damage, and no
sequoia to its present range and grove boundaries? How does fire and fire large Bigtree killed by them has ever been found."
frequency affect disease and insect interrelationships in the giant sequoia/
mixed-conifer ecosystem? Are current forest management strategies (e.g., It is finally being recognized that giant sequoia is
fire suppression, prescribed burning programs) affecting these interactions? subject to the same natural forces as other tree species (Bega
Giant sequoia trees are subject to the same natural forces (e.g., insect and 1964, Harvey and others 1980, Parmeter 1987, Piirto 1977,
disease organisms) as other tree species. An attempt is made in this paper to 1984b, Piirto and others 1974, 1977, 1984a, Weatherspoon
discuss some of the more common insect and disease associates of giant
sequoia and their significance in relation to the more complex temporal 1990). Weatherspoon (1990) has reported that "Although
(e.g., succession, aging and other time related events) and ecosystem inter- diseases are less troublesome for giant sequoia in its natural
relationships at work in the giant sequoia/mixed-conifer ecosystem. range than for most other trees, the species is not immune to
disease as once assumed." Scientists have been recording
disease associations with giant sequoia for some time
Giant sequoia (Sequoia gigantea [Lindl.] Decne.)3 may be (Seymour 1929, Bega 1964, Hepting 1971). But the signifi-
considered a long-lived species because of its high resistance cance, ecological role, and influences that affect these
to damage and mortality by insects, disease, and fire. Various organisms is not well understood.
attributes such as thick, fibrous bark, scalelike leaves, heart- Further, recent research on insects associated with
wood phenolic and tannin extractive content, general lack of giant sequoia has set aside many long-standing misleading
resinous extractives in bark and wood, and rapid growth rates generalizations about insect relationships in giant sequoia
of young trees may be partly responsible for enabling (David and others 1976, David and Wood 1980, DeLeon
individual giant sequoia trees to become very large and long- 1952, Stecker 1980a,b, Tilles 1984, Tilles and Wood 1982,
lived. We know very little, however, about what happens to 1986). Stecker (1980a) in this regard reports that "In sum-
giant sequoia throughout its lifetime. And, virtually nothing is mary, the giant sequoia, the largest living organism past or
known about predisposing growth-loss and mortality agents present, and one of the oldest, is unusual in having relatively
of young-growth giant sequoia in its native range. small insects comprising a relatively small insect fauna."
Many people assume that once giant sequoia seeds These research findings have caused some people still to
germinate, they will live a very long time and become very conclude that insect depredations are not seriously harming
large trees. It is not hard to understand why people still giant sequoias. Yet carpenter ants have been directly reported
believe this, since the older literature is replete with state- as being associated with old-growth giant sequoia trees (David
ments that giant sequoia has relatively few insect and disease and others 1976, David and Wood 1980, Piirto 1976, 1977,
pests. For example, the statement by John Muir (1894) Piirto and others 1984, Stecker 1980a, Tilles 1984, Tilles
illustrates this misconception: "I never saw a Bigtree [giant and Wood 1982, 1986). Carpenter ants, bark beetles, wood
sequoia] that had died a natural death; barring accidents they borers, and a variety of other insects are commonly found in
seem to be immortal, being exempt from all diseases that basal fire scars of old-growth giant sequoia. The role of
these insects in predisposing a giant sequoia tree to failure
or possibly in vectoring disease organisms is not completely
1 understood (David and others 1976, Piirto 1976, 1977, Piirto
An abbreviated version of this paper was presented at the Symposium
on Giant Sequoias: Their Place in the Ecosystem and Society, June 23-25, and others 1984). And, even though the insect fauna seems
1992, Visalia, California. to be smaller for giant sequoia based on present reports
2
Professor and Registered Professional Forester, Natural Resources (Stecker 1980a), it is inconclusive that insects are not a
Management Department. California Polytechnic State University, San Luis
problem to giant sequoia. The objectives, then, of this paper
Obispo, CA 93407
3
The correct scientific name for giant sequoia is currently a subject for are to: briefly review some of the more common disease and
disagreement. The common name, giant sequoia, and the scientific name insect organisms associated with giant sequoia; and discuss
Sequoia gigantea, will be used in this paper. Justification for this is detailed both temporal and ecosystem disease and insect interrelation-
in Davidson (1972) and in Piirto (1977). The common name, coast redwood,
ships at work in the giant sequoia/mixed-conifer ecosystem.
will refer to Sequoia sempervirens.

82 USDA Forest Service Gen. Tech. Rep.PSW-151. 1994

 Insect Relationships
(Sabuloides caberata and Pero behresarius), and
The scientific study by Stecker (1980a) is the most cutworms (Noctuidae spp.) (Parmeter 1987, Harvey 1980,
complete work to date on insects associated with giant Stecker 1980b, Metcalf 1948).
sequoia. Limited entomological research involving giant Very little scientific entomological research has been
sequoia occurred prior to 1974 (DeLeon 1952, Fry and White done to determine the ecological importance of some of the
1930, Hopkins 1903, Keen 1952, Person 1933). DeLeon insects listed by Stecker and others (1980a,b). Stecker (1980b)
(1952) was one of the first entomologists to dispute the determined that Phymatodes nitidus is very important in the
claim that coast redwood and giant sequoia have few insect continual release of seeds and subsequent reproduction of
enemies. DeLeon (1952) listed twenty insects which may giant sequoia. Researchers at the University of California at
use giant sequoia during all or part of their life cycles, Berkeley found carpenter ants (Camponotus modoc), which
including: bark and ambrosia beetles; flatheaded, roundheaded excavate nests in giant sequoia to rear their young, feeding
and other borers; leaf feeders; termites; carpenter bees; and on dead arthropods and honeydew from attended aphids
carpenter ants. (David and others 1976, David and Wood 1980, Tilles 1984,
Stecker (1980a,b) conducted detailed investigations on Tilles and Wood 1982, 1986). The attended aphid colonies
the invertebrates associated with giant sequoia seedlings, were found in the upper crowns of nearby white fir trees
living trees and fallen trees. Stecker's sample sites included (Abies concolor [Gord. and Glend.] Lindl. ex. Hildebr.).
Redwood Canyon, Redwood Mountain, Whitaker's Forest, This research finding has some very interesting implica-
Converse Basin, and Mountain Home State Forest. Stecker tions. For example, do the changes in white fir stand density
(1980a) found that: in giant sequoia groves (which have occurred as a result of
Of the 143 species of insects encountered in this fire suppression programs) cause associated changes in aphid
study, 4 species were found on seedlings (3 of which populations which in turn influence carpenter ant popula-
were also in older trees). Thirty-two insects were tions found nesting in living giant sequoia trees? Carpenter
found in different stages of downed and dead limbs ants are known to cause excavations that are so extensive as
of standing giant sequoias, while 3 of these were to seriously impair structural stability (Furniss and Carolin
also observed in standing dead wood of the living 1977, Piirto 1977, Piirto and others 1984b). Piirto and others
host. One hundred and fourteen insect species were (1984b) found that:
found only in the canopy of the living tree. Carpenter ants were found in or near the failure zone
Stecker (1980a) explains the results of his scientific in 16 [out of 33] study trees: only six of these,
investigations as follows: however, contained carpenter ant galleries in the
immediate failure zone. Microscopic examination
The insect fauna of the giant sequoia is small when of discolored wood associated with carpenter
compared to that of other conifers of the same ant galleries revealed an early to moderate
region. It differs considerably from that treated by stage of decay.
Southwood (1961) in the hypothesis that insect
species associated with a tree are a reflection of the A check of recent research records at Sequoia-Kings
cumulative abundance of that tree throughout Canyon National Park (Esperanza 1992) showed very little
recent geologic history. Insect faunas of histori- entomological research done since 1982 on insect associates
cally newer trees are considerably larger than those of giant sequoia. Piirto and others (1992a,b) identified six
of the much older giant sequoia. The insect faunas orders of insects found in association with giant sequoia fire
of pines and firs are usually two to three times scars. Thirteen individual insect species were identified, six
greater than the presently known fauna from the of which have not been reported previously. Insects identified
giant sequoia. most frequently were Ichneumonidae spp. (Ichneumons),
Camponotus spp. (carpenter ant), Curculionidae spp. (snout
Several of the insects listed in his study such as termites, beetle), Ceuthophilus spp. (cave cricket), and Blapylis alticolus
defoliators, various bark beetles (ambrosia beetle and B. productus (darkling beetles). On an infrequent basis,
Gnathotrichus sulcatus LeConte and Phloesinus punctatus seven insects were found associated with giant sequoia fire
LeConte), wood borers (Semanotus ligneus amplus Casey scars. These insects were identified as Sphecidae spp. (thread-
and Trachykele opulenta) and carpenter ants (Camponotus waisted wasp), Pompilidae spp. (spider wasps), Dorcasina
modoc) can be very destructive to young- and old-growth grossa (long-horned beetle), Cantharidae spp. (soldier beetle),
giant sequoia. T. opulenta larvae mine in the wood around Anthomyiidae spp (Anthomyid fly), Raphidiidae spp.
fire scars and apparently can also develop wholly in the bark (snakeflies), and Coreidae spp. (leaf-footed bug). More
(Fumiss and Carolin 1977). In addition, prominent insects research is needed, however, to determine the full ecological
associated with seedling damage and mortality include the significance of insects associated with giant sequoia both
camel cricket (Pristocauthophilus pacificus), two geometrids inside and outside its native range.

USDA Forest Service Gen. Tech. Rep.PSW-151. 1994. 83

Disease Relationships Southwest Research Station in Albany, California). These
lists are by no means all encompassing, as much remains to
For the purpose of this paper disease is defined as any be learned about disease relationships in giant sequoia.
deviation in the normal functioning of a plant caused by Comparatively little work has been done on the biotic
some type of persistent agent (Manion 1991). The abiotic and abiotic agents which damage seeds, seedlings, saplings
and biotic agents that cause disease are referred to as and young-growth giant sequoia trees. Cone and seed molds,
pathogens. It is important to recognize that diseases which damping off fungi and rootrot fungi are suspected as being
develop in plants and trees are the product of the tree/plant, major factors in preventing seedling establishment in native
the pathogen, and the interactions of these with the environ- giant sequoia stands (Stark 1968a,b; Parmeter 1987). These
ment over time. Time is an important factor as it is related fungi have been shown to cause reduced seed viability and
to both the rate at which a disease develops within an to prevent seedling establishment in undisturbed duff and
individual plant or tree and to the spread and increase of the litter areas within coast redwood stands (Davidson 1972).
pathogen population within a host population (Manion 1991). The role that these fungi play in native giant sequoia stands
Abiotic agents that can cause disease in forest trees is not known. Shellhammer and others (1971) reported that
include air pollution, high temperatures, freezing tempera- mice (Microtus spp.), gophers (Thomomys spp.), Cerambycid
tures, phytotoxic gases, pesticides, moisture stress, salt, poor beetles (Semanotus ligneus amplus) and a saprophytic
soil aeration, nutritional deficiencies/imbalances, mechanical fungus Hyphloma spp. possibly fasciculare were associated
damage and other abiotic factors. Any one or all of these with dead and dying 10- and 11-year-old giant sequoias
factors may affect giant sequoia at one time or another in the Abbot Creek drainage of the Cherry Gap Grove, a
during its life cycle. For instance, significant scientific previously logged area near Converse Basin.
information is becoming available on the impacts of air There are no known occurrences of any true mistletoe or
pollution on Sierra Nevada forests. Studies have been com- dwarf mistletoe species on either coast redwood or giant
pleted and many are underway which have/are evaluating the sequoia (Hawksworth 1978, 1992).
impacts of air pollution, climate change, acid rain and a
number of other abiotic agents which affect these forest Factors Associated With Tree Failure
ecosystems. Even though it seems apparent that abiotic agents
A number of major findings were reported in a study
like air pollution are having dramatic impacts on the giant
evaluating causes for tree failure of old-growth giant
sequoia/mixed-conifer ecosystem, giant sequoia trees are
sequoia trees (Piirto 1977, Piirto and others 1984a,b). Thirty-
rated as being most resistant to "smog" damage (Miller
three tree failures were evaluated in this study. Of the 33
1978). It is not known how this resistance varies with age of
failures listed, 21 percent apparently fell mainly because of
individual giant sequoia trees. The reader is referred to Manion
poor footing (wet soil, stream undercutting, etc.), 67 percent
(1991) for a general discussion of abiotic disease agents.
because of the failure of decayed roots and 12 percent
Some of the common biotic agents that cause disease in
because of stem breaks. All but two trees (both fell because
forest trees include nematodes, viruses, bacteria, fungi, and
of poor footing) had decay in either the stem or roots.
parasitic flowering plants (e.g., dwarf mistletoes). The
Carpenter ants were found in 16 trees but appeared to
symptoms of diseases caused by these biotic agents are
contribute to failure of only six. Fire scars were present on
usually expressed in disturbed or abnormal physiology of
27 trees and 26 fell to the fire-scarred side. This phenomenon
the host plant. Vascular wilt pathogens, for example, reduce
of giant sequoia tree failures falling to the scarred side was
the capacity of the vessels in trees to translocate water from
also reported by Hartesveldt and others (1975).
the roots to the top of the transpiring tree. These diseases are
typically classified into categories that are related to the Fungal Agents Associated With Decay and Tree Failure
stage of the host life cycle which the disease affects and/or
Many Basidiomycetes are responsible for the decay
to the expressed physiological effect seen in the forest tree.
observed in giant sequoia trees. Heterobasidion annosum,
Diseases caused by biotic agents include seedling diseases,
also called Fomes annosus, has been frequently observed in
leaf and needle diseases, root and trunk rots, shoot blights,
both the upper and lower stems of recent tree failures of
vascular wilts, or stem diseases. Full lists of these diseases
giant sequoia trees (Piirto 1977, 1984a,b, Piirto and others
associated with giant sequoia can be found in Seymour (1929),
1974,). Recent research (Piirto and others 1992a,b) is
USDA (1960), Bega (1964), Hepting (1971), Davidson (1972),
shedding new light on the hypothesis put forward by Piirto
Peterson and Smith (1975), Piirto (1977), Piirto and others
(1977) and Piirto and others (1984b) involving increasing
(1984a,b), and Parmeter (1987). Additional unpublished lists
stand density levels of white fir and other associated trees
of disease organisms associated with giant sequoia can also
species in giant sequoia groves. White fir is highly susceptible
be found in the records maintained by various herbariums
to a variety of forest diseases, particularly H. annosum.
(e.g., University of California at Berkeley, USDA Center for
Otrosina and others (1992) and Piirto and others (1992)
Forest Mycology Research at the Forest Products Laboratory
reported that both H. annosum isolates from white fir and
in Madison, Wisconsin and the USDA Forest Service Pacific
giant sequoia are of the ‘S’ intersterility Group meaning that

84 USDA Forest Service Gen. Tech. Rep.PSW-151. 1994

 they are interfertile. Given the interfertility of isolates Fire/Pathogen Interactions
collected from giant sequoia and true fir, an increase of Research recently completed by Piirto and others
white fir density in the absence of natural and prescribed fire (1992a,b) identified 17 new fungi from giant sequoia fire
may result in the build-up of H. annosum inoculum that scar wood samples. The fungi most frequently isolated were
could affect giant sequoia trees. H. annosum may spread along Byssochiamys fulva (a heat resistant fungus), which was
with other means via root contacts from white fir to giant isolated from 34 out of 90 fire scars sampled (38 percent);
sequoia. In greenhouse seedling inoculation studies, isolates Acrodontium intermissum (a preservative detoxifier),
of H. annosum collected from true fir and isolates collected isolated from 22 out of 90 fire scars (24 percent); and
on giant sequoia were capable of causing pathogenesis on Tritirachium sp. (an entomogenous fungus), which was
either species (Piirto and others 1992). isolated from 14 out of 90 fire scars (16 percent). Other
Other Basidiomycetes of particular note include Poria interesting fungi isolated include Neosartorya fischeri (a
albipellucida, Armillaria mellea, Poria incrassata, Stereum heat resistant fungus), Epicoccum nigrum (a soft rot fungus),
hirsutum, and others (Piirto 1977, Piirto and others 1984a,b). Leptographium sp. (L. wageneri causes black stain root
Armillaria spp. as receiving increasing attention as being an disease), Hyalorhin-ocladiella sp. (pentachlorophenol tol-
important pathogen of giant sequoia both within and outside erant; see Wang 1989), Mariannaea elegans (capable of
the native range of giant sequoia (Libby 1982). Libby (1992) forming soft rot cavities in European birch and Scots pine
reports Armillaria spp. as being the possible cause of wood; Levy 1969), and Basidiomycetes (fungi causing white
mortality in young-growth giant sequoia trees (up to 20 years or brown rots). A positive identification was made of one of
of age) planted in the Rhine River area of Germany. Trees the four confirmed Basidiomycete cultures isolated from
older than twenty years of age seemed to be less susceptible giant sequoia fire scars. Referred to as Phlebia subserialis
to Armillaria spp. and hardwoods were known to have [(Bourd. et Galzin) Donk] of the Corticiaceae family and
occurred in this Rhine River giant sequoia plantation. known to affect both hardwoods and softwoods this organism
Another recent and interesting report from Libby (1992) is classified as a white rotter which may have potential in
involves the forty-year old plantation of giant sequoia trees biological pulping processes (Dorworth 1992, Burdsall 1992).
around the Forest Hill Seed Orchard. The study showed that The majority of these fungi identified in the Piirto and
up to forty percent of the planted young giant sequoia trees others study (1992a,b) are microfungi. As described by
which are between 30 and 50 feet tall and up to 10 inches in Wang and Zabel (1990), microfungi are ubiquitous and
diameter at breast height had thin crowns and bunches of cosmopolitan. Some are plant pathogens and some are human
trees were easily pushed over. Preliminary observations by pathogens, but most are saprobes that derive nourishment
Forest Service pathologists present with Bill Libby at from decaying organic matter that occupy all segments of
the Forest Hill site indicate that Armillaria spp. may be the environment, most commonly the soil. The term
responsible for the mortality that is occurring there. In the microfungi is commonly used to describe fungi belonging to
Forest Hill situation, stumps of California black oak (Quercus the Subdivision Ascomycotina. These microfungi have been
kelloggii Newb.) and dead deer brush (Ceanothus integerrimus found associated with utility poles made of Douglas fir,
H.&A.) were in the immediate vicinity of the dead giant southern pine, and western red cedar. Some have shown to
sequoia trees. It can be inferred from previous research and cause soft rot and degradation of wood structure. Recent
from the Rhine River and Forest Hill Seed Orchard research suggests some of the microfungi can also reduce
situations that 1) Armillaria spp in combination with other the fungitoxicity of wood preservative chemicals (Wang and
organisms may be important pathogens of both young- and Zabel 1990). Based on previous research, it is possible the
old-growth giant sequoia; 2) Armillaria spp seems to be microfungi isolated from giant sequoia may play a role in
particularly aggressive in areas where hardwoods and the decay process. An interesting organism for instance, that
possibly vulnerable species of Ceanothus spp. occur; 3) giant has been known for some time to be frequently found on
sequoia trees planted outside their natural range may be exudate of exposed heartwood of coast redwood and giant
particularly vulnerable to a whole host of disease problems sequoia fire scars is Mycocallicium sequoiae (Bonar 1971,
which as of yet are not completely understood. Piirto 1977, Piirto and others 1992a). More research is needed
Bega (1964), Libby (1982), Worral and others (1986) on the variety of microorganisms at work in the giant
have conducted studies on giant sequoia trees planted sequoia/mixed-conifer ecosystem.
outside their native range and found some of them severely Microorganisms and pathogens can be affected in a
damaged by a canker fungus Botryosphaeria sp. In addition, variety of ways by fire; and while none of these effects has
giant sequoia seedlings in nurseries were found to be been adequately evaluated for giant sequoia ecosystems,
particularly vulnerable to charcoal root disease caused by some possible fire effects with important management
Macrophomina phaseoli and grey mold caused by Botrytis implications have been described (Parmeter 1977, Parmeter
cinerea (Peterson and Smith 1975). Understanding the and Uhrenholdt 1975, 1976, Piirto 1977). It has been
organisms that cause disease and decay in giant sequoia is discussed that if Heterobasidion annosum may build up on
important, but it is equally important to understand the inter- white firs (Abies concolor) and then attack nearby giant
actions that occur between organisms and the environment. sequoias (Piirto 1977), fires may "cauterize" giant sequoia

USDA Forest Service Gen. Tech. Rep.PSW-151. 1994. 85

fire scars and thus reduce decay (Christensen and others species and environmental conditions. This has been empha-
1987). While these aspects are recognized and in need of sized by Hepting and Shigo (1972) for oaks in North
study, it is likely other fire effects on both the host and Carolina and Maine. It is equally important to know that
associated pathogens occur and only research monitoring once infection has occurred, survival and damage may
these fire effects on microorganisms will allow recognition continue throughout the life span of a tree. This has been
and further evaluation. These fire/pathogen interactions in reported for fire-damaged white fir (Abies concolor)
the giant sequoia/mixed-conifer ecosystem are reported in regeneration (Aho and Filip 1982).
Piirto and others (1992a,b). In a recent study conducted by Piirto and others (1992a,b),
The management implications of how a disease impacts it was determined, based on a quantitative evaluation of 90
stands of giant sequoias is incomplete (Parmeter 1987). living old-growth giant sequoias, that as much as 70 percent
Knowledge of giant sequoia disease is increasing, but specific of the circumference (values ranged from 3.3 percent to 69.5
effects on regeneration, stand development, old-growth and percent with an average of 27.3 percent) and as much as 54
young-growth trees are still relatively unknown. percent of the cross-sectional area (values ranged from 3.2
percent to 53.7 percent with an average of 15.6 percent)
Wounds As Entrance Courts were adversely affected by fire scars at groundline. This
research work further demonstrated that prescribed burning
Following fire, trees may show reduced growth and programs caused damage to callus tissue around existing
vigor, owing mainly to heat injury (Hare 1961) or perhaps to fire scars 52 percent of the time, with recessional and
"shock" if stand density is changed. The ecological effects enlargement damage noted 57 percent and 35 percent of the
of this damage can result in numerous pathogens and insects time respectively.
adapted to attack the weakened yet surviving plants. The The role of fire scars on giant sequoia trees as entrance
literature is replete with reports noting a strong association courts for decay and insect attack has been defined on the
of insect, pathogen, and microorganism activity in fire basis of numerous research studies. As we move forward
damaged trees. In all of these reports, fire damaged roots with the reintroduction of fire in the form of prescribed
and/or above ground basal fire scars served as an infection burns in giant sequoia grove areas, the questions still remain
court for these organisms. as to what intensity, where, when, and how much should be
Perhaps the best documented effects of fire on disease burned to largely promote the positive values fire provides to
involve the creation of infection courts for heart rot fungi the giant sequoia/mixed-conifer ecosystem while minimizing
(Boyce 1961; Harvey and others 1976; Hepting 1935; Nelson the negative consequences to specimen old-growth giant
and others 1933; Nordin 1958). Heart rot which frequently sequoia trees. The results by Piirto and others (1992a,b)
develops in fire scars may in turn effect other processes, confirm that the valued older trees in the giant-sequoia/
such as bark beetle activities in burned stands (Geiszler and mixed-conifer system are damaged, and that fire management
others 1980a,b). Canker fungi may also be associated with strategies should be implemented to minimize these adverse
fire scars (Hinds and Krebill 1975). Decay following fire fire impacts.
scarring can reduce productivity in timber stands. It can also
lead to hazard problems and to the loss of valuable specimen Summary and Conclusions
trees in parks and preserves, as may be the case with loss of
giant sequoias (Piirto 1977). Gill (1974) reported secondary It is important to consider several factors when
effects of fire on fire scars often lead to mechanical failure interpreting insect and disease relationships in the giant
of Eucalyptus pauciflora. sequoia/mixed-conifer ecosystem including:
Perry and others (1985) report that damage to living
jarrah (Eucalyptus marginata), Karri (E. diversicolor), and 1. No single variable functions alone because insects,
maritime or cluster pine (Pinus pinaster) in southwestern disease organisms and the ecosystems operate
Australia by fire or mechanical injury during forest opera- interdependently.
tions emerges as a major factor facilitating the entry of fungi 2. Various microorganisms have beneficial and antago-
and termites that spread and degrade substantial areas of nistic influences on disease evelopment. Hepting
heartwood. Littke and Gara (1986) and Gara and others (1935) was one of the first forest pathologists to
(1986) report that the extension of decay columns from fire note an association of various microorganisms (e.g.,
damaged roots up into the boles of lodgepole pine (Pinus bacteria, Trichoderma spp. and Penicillium spp.)
contorta var. murrayana) suggests root damage is their most with decay fungi. These organisms can inhibit or
important source of stem decay. Newly fire-damaged root facilitate decay development in forest trees. Many
tissues were infected with imperfect fungi and white rot such known and unknown interactions are at work
basidiomycetes. Decayed material from previous fires yielded in the giant sequoia/mixed-conifer ecosystem.
primarily brown rot basidiomycetes. It is important to know 3. Fire does influence the type and population levels of
that the amount of microorganism, pathogen and insect a vast variety and number of insects, pathogens and
activity associated with fire scars is influenced by the tree microorganisms found in the giant sequoia/mixed-

86 USDA Forest Service Gen. Tech. Rep.PSW-151. 1994

 conifer ecosystem. These interactions are not well sequoia. I also want to thank the USDA Forest Service for
understood. These relationships are further compli- allowing me to work on an intermittent basis for the Sierra
cated by changing climatic patterns such as drought. and Sequoia National Forests. This work experience has
For example, a bark beetle like Phloesinus punctatus enabled me to keep close to the issues that are confronting
(western cedar bark beetle), which is common in this nation with reference to the management of our public
fallen branches of giant sequoia could, in drought lands. Grateful appreciation is extended to Mr. Kevin Piper,
periods, become more detrimental (Wood 1992). Ms. Amy Workinger, Dr. John R. Parmeter Jr., Dr. David L.
More research is needed. Wood, Dr. Fields Cobb Jr., Dr. June Wang, Dr. Harold
4. Further complicating these complex ecosystem Burdsall, Dr. W. Wayne Wilcox, Dr. Tom Chase, Dr. William
interactions is the influence of human activities. Otrosina, Dr. David Parsons, Dr. Jan van Wagtendonk and
Fire suppression, whereas beneficial in many ways, many others too numerous to list here for their research
has altered stand development in the giant sequoia/ assistance over the last several years.
mixed-conifer ecosystem. Changes in stand density
and species makeup may be aggravating disease
and insect relationships (Piirto 1977). Prescribed References
burning, on the other hand, has both beneficial and
detrimental effects. Wounds caused by fire have Aho, P.E. ; Filip, G.M. 1982. Incidence of wounding and Echinodontium
unquestionably been associated with both disease tinctorium infection in advanced white fir regeneration. Canadian
Journal of Forest Research 12(3): 705-708.
and insect attack in forest trees (Manion 1991). Bega, R.V. 1964. Diseases of sequoia. In: Diseases of widely planted
5. Young-growth giant sequoia both in and planted forest trees. FAO/IUFRO Symposium on internationally dangerous
outside its native range are vulnerable to a whole forest diseases and insects; Oxford: 1964 July 20-30, 131-139.
host of insect and disease problems. Botryosphaeria Bonar, L. 1971. A new Mvcocalicium on scarred sequoia in California.
Madrono 21(2): 62-69.
spp and Armillaria spp appear to be two major Boyce, J.S. 1961. Forest Pathology. New York: McGraw-Hill Book Co.;
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