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Wildlife, Animals, and Plants
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Introductory
SPECIES: Lithocarpus densiflora | Tanoak
ABBREVIATION :
LITDEN
SYNONYMS :
Lithocarpus densiflorus
Quercus densiflora
SCS PLANT CODE :
LIDE3
COMMON NAMES :
tanoak
tanbark-oak
tan oak
TAXONOMY :
The currently accepted scientific name of tanoak is Lithocarpus
densiflora (Hook. & Arn.) Rehd. [17,34].
Tanoak is the only North American representative of a large Asian genus.
The genus Lithocarpus is considered a link between true oaks (Quercus)
and the chinquapins and chestnuts (Castanopsis and Castanea), possessing
characteristics of each [34,42]. A shrubby growth form of tanoak,
commonly known as shrub tanoak, has been recognized at the varietal
level as Lithocarpus densiflora var. echinoides (R. Br.) Ambrams.
[17,34]. Unless stated otherwise, this discussion pertains to the
typical variety L. densiflora var. densiflora.
LIFE FORM :
Tree
FEDERAL LEGAL STATUS :
No special status
OTHER STATUS :
NO-ENTRY
COMPILED BY AND DATE :
N. McMurray, June 1989
LAST REVISED BY AND DATE :
N. McMurray, August 1989
AUTHORSHIP AND CITATION :
McMurray, Nancy E. 1989. Lithocarpus densiflora. In: Remainder of Citation
DISTRIBUTION AND OCCURRENCE
SPECIES: Lithocarpus densiflora | Tanoak
GENERAL DISTRIBUTION :
A major component of coastal mixed evergreen forests, tanoak is
distributed from the Cascade Mountains of southwestern Oregon southward
through the Klamath Mountains and California Coast Ranges to Ventura
County [25,34,42]. In northern California, tanoak ranges inland to the
lower slopes of Mount Shasta and occurs intermittently along the west
slope of the Sierra Nevada as far south as Mariposa County [31]. Sierra
stands are locally abundant in Butte and Yuba Counties [13]. Recent
inventories indicate that this hardwood comprises the dominant cover
type over at least 861,000 acres (350,700 ha) in California [4].
Shrub tanoak (var. echinoides) occurs in chaparral communities
throughout the mountains of northern California and southern Oregon,
becoming particularly abundant in the vicinity of Mount Shasta
[31,34,42]. Scattered populations also occur in the southern Coast
Ranges and in the central Sierra Nevada [34].
ECOSYSTEMS :
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES23 Fir - spruce
FRES26 Lodgepole pine
FRES27 Redwood
FRES28 Western hardwoods
FRES34 Chaparral - mountain shrub
STATES :
CA OR
ADMINISTRATIVE UNITS :
REDW YOSE
BLM PHYSIOGRAPHIC REGIONS :
1 Northern Pacific Border
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains
KUCHLER PLANT ASSOCIATIONS :
K004 Fir - hemlock forest
K005 Mixed conifer forest
K006 Redwood forest
K007 Red fir forest
K026 Oregon oakwoods
K028 Mosaic of K002 and K026
K029 California mixed evergreen forest
K030 California oakwoods
K033 Chaparral
SAF COVER TYPES :
207 Red fir
211 White fir
229 Pacific Douglas-fir
230 Douglas-fir - western hemlock
231 Port Orford-cedar
232 Redwood
233 Oregon white oak
234 Douglas-fir - tanoak - Pacific madrone
243 Sierra Nevada mixed conifer
244 Pacific ponderosa pine - Douglas-fir
245 Pacific ponderoas pine
246 California black oak
247 Jeffery pine
248 Knobcone pine
249 Canyon live oak
255 California coast live oak
SRM (RANGELAND) COVER TYPES :
NO-ENTRY
HABITAT TYPES AND PLANT COMMUNITIES :
Self-perpetuating stands of tanoak are indicative of climax conditions
in a number of communities within evergreen hardwood [35], mixed
evergreen [1,10,55], redwood (Sequoia sempervirens) [47,55], and mixed
conifer forests [44].
Within mixed evergreen forests in southwestern Oregon, Atzet [1]
describes climax tanoak communities associated with warm, moist sites
along the lower slopes of the Siskiyou Mountains. Even though most
stands are currently dominated by a Douglas-fir (Pseudotsuga menziesii)
overstory, Douglas-fir is a fire-maintained, seral component within
these stands. Climax understory dominants within the tanoak series may
include vine maple (Acer circinatum), Cascade holly grape (Berberis
nervosa), poison oak (Rhus diversiloba), salal (Gaultheria shallon),
vanilla leaf (Achlys triphylla), common princes pine (Chimaphilla
umbellata), and twinflower (Linnaea borealis).
VALUE AND USE
SPECIES: Lithocarpus densiflora | Tanoak
WOOD PRODUCTS VALUE :
Tanoak compares favorably with eastern hardwoods in strength, hardness,
and machining characteristics [31]. Manufacture of tanoak wood products
has been limited, however, due to an inconsistent supply of quality raw
materials. When processed properly, upper grades exhibit excellent
strength and an oaklike appearance, and produce a good grade of veneer
and plywood. Tanoak flooring, panelling, and decking have also been
manufactured [43]. Where stength, not appearance, is of primary concern,
lower grades have been used to make pallets, crossties, mine timbers,
baseball bats, and garden tool handles [7,31,43]. The wood has also
been chipped for particle board and pulp manufacture. Besides being
widely used as a source of domestic fuelwood [7], tanoak is currently
under consideration for use in waferboard, as a sports equipment
laminate, and for the cogeneration of electricity [31].
IMPORTANCE TO LIVESTOCK AND WILDLIFE :
Livestock: Although tanoak is generally considered of little browse
value to the majority of livestock [6,45], acorns and young seedlings
are highly preferred by hogs and cattle [43]. The leaves of shrub
tanoak (var. echinoides) provide some forage for domestic goats; use
typically corresponds to a lack of other more palatable forage and is an
indication of overgrazing on many sites [45].
Wildlife: Tanoak provides important habitat for numerous wildlife
species [2]. Because it tends to grow in mixtures with hardwoods and
conifers, tanoak is often a component of communities which are
structurally and compositionally diverse [31]. Tanoak habitats supply
food and nesting sites for the northern flying squirrel, Allen's
chipmunk, and dusky-footed woodrat [39]. Mammals which feed on tanoak
acorns include the black bear, black-tailed deer, Townsend chipmunk,
California ground squirrel, and redwood chickaree. Acorns are also a
source of food for birds such as the band-tailed pigeon, California
woodpecker, and varied thrush [39,42,53]. Cavity nesters known to use
tanoak include the downy woodpecker, northern flicker, red-breasted
nuthatch, white-breasted nuthatch, brown creeper, and house wren [40].
PALATABILITY :
Tanoak is of low palatability to most livestock and wildlife [6,45].
Utilization occurs primarily when other more preferred species are
unavailable. Mule deer readily consume both the leaves and acorns of
shrub tanoak (var. echinoides) in Yosemite National Park [45].
NUTRITIONAL VALUE :
Nutritional content of acorns produced by a number of West Coast
hardwoods including tanoak, California black oak (Quercus kelloggii),
valley oak (Q. lobata), blue oak (Q. douglasii), interior live oak (Q.
wislizenii), canyon live oak (Q. chrysolepis), and California live oak
(Q. agrifolia) are presented below [24].
carbohydrate 42 - 52%
protein 3 - 5%
fat 4 - 14%
COVER VALUE :
The multilayered structure of mature tanoak stands provides hiding cover
for a variety of birds and small mammals [39]. The northern flying
squirrel, Allen's chipmunk, and dusky-footed woodrat all rely heavily on
tanoak habitats for hiding, thermal, and nesting cover. Tanoak logs are
utilized as resting and hiding cover by a number of salamanders
including the ensatina, Del Norte salamander, and black salamander.
VALUE FOR REHABILITATION OF DISTURBED SITES :
Tanoak may be useful for controlling erosion since plants reestablish
rapidly through vigorous sprouting following disturbance [43].
OTHER USES AND VALUES :
Tannin produced from tanoak bark is used commercially to cure leathers
[31,42,53]; the tannin is particularly well suited for the curing of
heavy leathers such as soles of shoes and saddles [42]. Tanoak tannin
also possesses some medicinal properties. Ground acorns have been used
as chicken feed. Tanoaks are occasionally cultivated as ornamentals
[23]. The current year's growth exhibits a fuzzy covering of
reddish-brown hairs which often irritates the eyes and skin and may
cause coughing and sneezing [4,42].
Historically, tanoak acorns provided a dietary staple for indigenous
peoples throughout the California Coast Ranges. The tannins were
leached out, and then the acorns were either pounded into a mush or
dried and ground into flour for baking [22,43]. Apparently tanoak
acorns were much preferred over other types due to their high oil
content [42].
MANAGEMENT CONSIDERATIONS :
Competition: Tanoak represents a significant source of competition to
conifers [4,29,38]. Even though plants do not resprout as aggressively
as many associated species such as bigleaf maple (Acer macrophyllum),
madrone (Arbutus menzeisii), or Oregon white oak (Quercus garryana),
tanoak is much more abundant in conifer understories [21,42,48]. After
release from heavy shade, suppressed tanoak develops with remarkable
speed, often forming a dense cover [31]. Prolonged tanoak competition
typically results in lost conifer growth, extended rotations, and
inadequate conifer stocking on many sites [31]. Tanoak competition may
be particularly severe in new plantations, causing slow growth and high
conifer seedling mortality. To maintain plantations, tanoak must be
suppressed to the point where conifers can gain dominance. Herbicides
can be effective in temporarily reducing tanoak [5,8,12,20]. Possible
nonchemical control methods include mechanical means of site preparation
such as uprooting stumps or stump grinding [32]. Preharvest burning or
brush slashing used in conjunction with either preharvest or postharvest
burning are also potential methods of control [15,18,26]. Nonchemical
treatments are most successful when applied within a few years of
planting. Postplanting, follow-up treatments may be necessary on many
sites. Biotic control agents have received some attention. The
uncommon development of stunted and chlorotic tanoak sprouts on logged
areas in northern California could not be linked to either pathogens or
viruses [32].
Prediction equations have been developed which reliably estimate the
postdisturbance sprouting potential of tanoak from preharvest inventory
data [48]. Since these equations predict cover by age group and
diameter class, it may be possible to identify that segment of the
tanoak understory contributing most to postdisturbance coverages,
thereby aiding in the selection of an appropriate control method on a
site by site basis.
Tanoak management: Tanoak has often been considered a "weed" tree with
respect to conifer management. Since it is a highly persistant species
on many sites, intensive management of tanoak for hardwood timber
production may be a logical management option on some sites [28,31].
Management potential is highest on sites where extensive, even-aged
stands have developed following clearcutting or fire. Clearcut
harvesting and manipulation of subsequent sprout stands is the
recommended silvicultural treatment [30]. Sprouting from burls may be
encouraged by cutting stumps to less than 8 inches (20 cm); basal
sprouts have a low incidence of rot bridging from stump to sprout pith.
Leaving higher stumps encourages stool sprouts and increases the
likelihood of heart rot in young stands [30,31]. Tanoak grows better
and with better form if crowns are all in codominant position. As the
number of sprouts per clump declines over time, stem density becomes a
significant factor in maintaining the mutual shading so necessary for
optimal growth. Consequently, best development tends to occur where
stands are composed of both high densities of tanoak sprout clumps and
multiple stems per clump [31]. Initial thinnings are most effective
after the age of 20, at which time growth has been concentrated into a
few dominant stems; thinning prior to this time causes the production of
numerous resprouts. In northern California, best growth is
obtained when thrifty, 20-year-old stands are thinned to basal areas of
approximately 90 square ft/acre (21 square m/ha). McDonald and
Tappeiner [31] suggest interplanting thinned stands of tanoak with
Douglas-fir. Tanoak could then be harvested earlier in the rotation,
thereby increasing the growth and yield of Douglas-fir.
BOTANICAL AND ECOLOGICAL CHARACTERISTICS
SPECIES: Lithocarpus densiflora | Tanoak
GENERAL BOTANICAL CHARACTERISTICS :
Tanoak is a slow-growing, shade-tolerant, native, sclerophyllous,
evergreen hardwood tree [31,33,42,53]. Tanoak ranges from 65 to 150
feet (20-45 m) in height [33], but mature trees usually do not exceed 82
feet (25 m) in height and 3.2 feet (1 m) in diameter. Tanoak may live
to be 300 to 400 years old; average age on better sites in the Pacific
Northwest is approximately 180 years [42]. The reddish-tinged, pale
brown bark becomes thick and fissured as trees mature [22,34,37]. Large,
leathery, blunt-toothed leaves are retained for approximately 4 years
[49]. Male and female flowers are borne in the leaf axils of new
shoots. The fruit is a large acorn approximately 1 to 2 inches (2.5-5
cm) long and 0.6 to 0.7 inch (1.5-1.8 cm) in diameter and usually borne
singly or in clusters of two to three [42]. At the soil surface, tanoak
possesses a woody, underground regenerative organ known as a lignotuber
[16,31,50]. The extensive root system consists of a deep taproot which
is associated with nitrogen-fixing bacteria [33].
Tanoak growth forms vary according to stand structure, stand
composition, and site quality. When occurring as part of the conifer
canopy, tanoak develops a long, clear central bole and an excurrent
branching habit similar to that of conifers. Where development beneath
conifers is delayed, understory tanoak may be multistemmed and/or
shrublike [42]. In more open hardwood stands, however, the main trunk
is usually short and thick and divides into several large, horizontal
limbs, forming a broad, rounded crown. Shrub tanoak (var. echinoides)
is a stunted form typically associated with chaparral vegetation.
Heights range from 1 to 10 feet (up to 3 m); the branches are rigid with
small, thin leaves [34,42].
RAUNKIAER LIFE FORM :
var. densiflora
Undisturbed State: Phanerophyte (mesophanerophyte)
Undisturbed State: Cryptophyte (geophyte)
Burned or Clipped State: Cryptophyte (geophyte)
var. echinoides
Undisturbed State: Phanerophyte (microphanerophyte)
Undisturbed State: Phanerophyte (nanophanerophyte)
Burned or Clipped State: Cryptophyte (geophyte)
REGENERATION PROCESSES :
Tanoak regenerates sexually and vegetatively. Seedling regeneration is
characterized by very slow recruitment rates. Once established,
however, tanoak individuals are extremely persistent beneath conifer and
hardwood overstories, maintaining themselves through sporatic stem
mortality and sprout regeneration. After being released from heavy
shade, suppressed tanoak "seedling banks" develop rapidly, often
dominating early successional stages.
Reproduction from seed: Tanoak sprout clumps can produce acorns as
early as 5 years of age; consistent and abundant production of viable
acorns usually occurs when trees are 30 to 40 years old [31,42]. Acorn
production is frequent and prolific with large seed crops produced
almost every other year [30,42]. Tanoak produced bumper seed crops four
times over a 24-year-period on sites in California [27]. Complete seed
failures are rare and typically coincide with drought or frost [42].
Acorns are dispersed in the fall of their second year and are up to 79
percent sound. Insect-infested acorns usually drop first. Most fall
directly beneath the parent plant although long-distance disperal may
occur via mammal and bird vectors [30]. Despite abundant acorn
production, tanoak reproduces extremely slowly via seed. Unless acorns
are rapidly covered by litter and leaves, the majority of acorns are
consumed by rodents, birds, or deer.
Germination rates vary from 19 to 80 percent; no dormancy-breaking
treatment is necessary [42]. On fall planted sites in southwestern
Oregon, tanoak germination and radical elongation began in late January
and early February; however, shoot emergence was delayed until mid July
[53]. Germination occurs much more rapidly in acorns positioned
point-up; alternate positions extend the germination period [27,31]. Some
germination also may occur during mild fall weather [42]. Whereas
optimal seedbeds consist of loose, moist mineral soil, tanoak also
establishes abundantly in deep litter [30,42,54]. Litter helps keep
acorns moist and provides insulation from temperature extremes. A deep
covering of litter, however, tends to hinder germination [42]. Acorn
viability is usually short-lived [31].
Successful seedling establishment occurs most frequently in the duff and
litter beneath existing conifer and hardwood stands [50,54]. A firm,
woody hypocotyl greatly lessens first year mortality caused by
litterfall [50]. Seedling growth is best in partial shade. Most early
growth is devoted to the formation of a lengthy taproot and shoot growth
is extremely slow by comparison. Four-year-old seedlings typically
average less than 6 inches (15 cm) in height [30]. Seedling
establishment is quite rare on disturbed sites [31]. Inadequate moisture
and an unfavorable environment generally contribute to poor seedling
survival on shrub-free clearcuts in northern California [27]. Initial
establishment success also appears to be related to differences in seed
predation on clearcuts versus forested habitats. Seed predation is
likely to be considerably higher on clearcut areas where small mammal
populations, particularly deer mice, are quite high [50].
Interestingly, survival and growth of emerged tanoak seedlings was
approximately equal over 4 years on clearcut and forested sites in
Oregon.
Vegetative regeneration: Since regeneration occurs in dry, low light
environments, tanoak maintains itself through periodic diebacks.
Frequent stem mortality and resprouting apparently enhance its
persistence on a site by reducing respiratory losses from inefficient,
senescing stems [49]. If left undisturbed, tanoak seedlings consist of
single stem until the 5 to 12 years of age. At this time the stem dies
back to the ground, a rudimentary underground burl develops at the stem
base, and from one to five resprouts are produced from perennating buds
located on the burl [31]. Burls enhance the sprouting potential of
tanoak by functioning as a source of numerous dormant buds [14,31] and
perhaps as storage for carbohydrates, nutrients, and water [16,31]. Burl
size increases with age. Tanoak burls are usually well-developed by age
60, sometimes reaching up to 4.8 inches (12 cm) in diameter. Burl
growth is related to site quality and is predictable on a given site
[31]. Bud numbers increase as burls enlarge; not only do preformed buds
divide, but adventitious buds also arise from callus tissues formed at
the base of dead stems. Therefore, sprouting potential increases as
tanoak grows. Some large tanoak stumps may have as many as 1,400 basal
buds (burl and stump surfaces) [42].
Young tanoak typically develop a multistemmed, shrublike growth form
beneath conifer and hardwood overstories. Stem mortality and
resprouting commonly occur at least 3 to 5 times before the age of 60
[49]. Sprouts which develop from these diebacks, in addition to those
after the initial dieback, are generally referred to as "seedling
sprouts". In young conifer-tanoak stands with high initial conifer
stocking, nearly half of the tanoak regeneration may loose all stems as
a result of conifer crown closure [49]. Stem mortality is most likely
in stems less than 6 inches (15 cm) in d.b.h. From 60 to 90 percent of
the tanoak sprouts less than 6 inches (15 cm) in d.b.h. may die in
younger stands. As a component of more mature conifer stands, however,
only 10 percent of the sprouts in this size class die.
Basal sprouting is a common characteristic of large, undisturbed tanoak
growing in older conifer stands [49]. Sprouts are frequently prostrate
and may occassionally layer. Most large tanoak have more than one burl.
One of the largest tanoak within a mature conifer stand in Oregon had
five stems ranging from 25 to 63 feet (7.6-19.3 m) in height and from 39
to 94 years in age. Twenty-two resprouts arose from the base.
Belowground, a rootlike structure connected four burls which ranged from
15.2 to 33.2 inches (38-83 cm) in diameter.
Tanoak also resprouts following fire or cutting [31,42]. Burning
younger, thin-barked tanoak generally kills aboveground portions of the
stem. As a result, basal sprouting is the primary means of postburn
regeneration. Following cutting, however, tanoak may also develop
"stool sprouts" near the top of higher stumps; stool sprouts originate
from either the vertical surface of the stump or from the cambium on the
horizontal cut [30,31]. Resprouting abilities develop exceptionally early
in tanoak. Following a frost which killed the aerial shoots of
1-week-old seedlings, 75 percent resprouted; most had two to four
resprouts per plant [31].
SITE CHARACTERISTICS :
Although adapted to a variety of sites [31], tanoak occurs most
abundantly along the lower slopes of coastal mountain ranges on sites
characterized by high levels of moisture and mild temperatures [1,34].
In southern Oregon and northern California where tanoak reaches its
maximum development, trees grow on east and north slopes at elevations
between 500 and 3,000 feet (152 and 915 m). Toward the southern portion
of its distribution, tanoak occupies westerly aspects between 2,400 and
4,700 feet (732 and 1,434 m) [31]. Tanoak also competes well inland,
becoming locally abundant along the west slope of the Sierra at
elevations between 1,900 and 5,000 feet (580 and 1,525 m). In warmer
environments of the central Sierra, tanoak typically occupies sites
having more favorable moisture regimes such as foothill riparian areas,
sheltered coves and ravines [42]. A ubiquitous species througout its
range, tanoak is commonly distributed as scattered individuals and in
clumps or groves within undisturbed conifer forests [31]. Extensive
stands often develop following logging or fire [31,42,36].
Climate: Annual precipitation ranges between 40 and 100 inches (102 and
254 cm), 70 percent of which falls between November and February [42].
Monthly precipitation during the dry summer and early fall is often less
than 1 inch (2.5 cm). Summer moisture is vital for tanoak survival [1].
Optimal sites are often associated with atmospheric moisture,
experiencing high humidities, summer fog, and low-lying clouds; such
conditions help to ameliorate temperature extremes. Generally, tanoak
is restricted by moisture limitations on shallow soils, and by low
temperatures at higher elevations [1].
Soils: Tanoak is most often associated with deep, fertile, well-drained
loamy, sandy or gravelly soils [31,42]. Its presence can be used as an
indicator of productive sites in southwestern Oregon [1]. Trees can
also grow well on stony or shallow soil sites located on north slopes.
Associates: As a component of hardwood, mixed hardwood and conifer
forests, tanoak is associated with a wide variety of species.
Douglas-fir and Pacific madrone are by far the most common tree
associates. Other common associates include [1,42]:
conifers - redwood, white fir (Abies concolor), sugar pine (Pinus lambertiana),
Pacific yew (Taxus brevifolia), and ponderosa pine (Pinus ponderosa).
hardwoods - canyon live oak (Quercus chrysolepis), California black oak,
interior live oak, California laurel (Umbellularia californica), and
giant chinquapin (Castanopsis chrysophylla).
shrubs - California hazel (Corylus rostrata var. californica), poison
oak, salal, pinemat manzanita (Arctostaphylos nevadensis); and Cascade
holly grape.
forbs - Torrey peavine (Lathyrus torreyi), and mottleleaf ginger (Asarum
hartwegi).
ferns - sword fern (Polystichum munitum) and bracken (Pteridium
aquilinum).
grasses - bromes (Bromus spp.) and fescues (Festuca spp.).
Shrub tanoak (var. echinoides): The shrubby form of tanoak is commonly
associated with chaparral communities. Plants occupy mountain tops and
exposed ridges at elevations between 2,000 and 8,000 feet (610 and 2,440
m). Soils are dry and rocky [34,42,36]. Shrub tanoak is often the
dominant form on serpentine soils [42].
SUCCESSIONAL STATUS :
Tanoak is a long-lived species capable of establishing beneath a full
canopy of hardwoods or conifers; establishment occurs under both
overstory and understory trees [42,54]. Substantial seedling
establishment is often delayed 25 to 30 years in conifer stands [49].
Once established, tanoak is extremely persistent on a site despite low
light levels. Whereas tanoak may attain tree size where gaps develop in
the overstory, suppressed tanoak individuals are shrublike, maintaining
themselves through periodic diebacks [31,42]. Prolonged periods without
disturbance permits the slow, steady, annual accumulation of seedlings.
Tanoak is able to survive disturbances such as fire, logging, blow-down,
insect devastation, or mass soil movement [10,42,53]. Plants sprout
vigorously once released from dense shade and are extremely competitive
[1,30]. Even-aged sprout stands develop rapidly from banks of suppressed
individuals and may dominate early seral situations.
Although tanoak may form almost pure, climax stands within evergreen,
hardwood forests [1,10,42], it more often occurs as part of the climax
subcanopy on sites within mixed evergreen, redwood, Douglas-fir, and
mixed-conifer forests [1,10,47,54,55]. If left undisturbed, tanoak
eventually forms uneven-aged stands [31,54]. Fire exclusion has favored
the establishment of tanoak within redwood and mixed evergreen forests.
On many sites, fire-maintained Douglas-fir are now being replaced by
tanoak and other more shade-tolerant species [1,54]. With continued
fire exclusion, tanoak is the potential climax dominant throughout much
of the mixed evergreen zone [1,10].
SEASONAL DEVELOPMENT :
Tanoak typically flowers from June through August, although blooming may
also occur during the spring or fall [34,42]. The foliage is often
concealed by the profuse, yellowish blooms. Apparently a long dry
period is essential for setting of acorns. Acorns ripen in the fall of
their second season and are generally dropped between September 20 and
November 15 [42].
Limited observations indicate the following elevational pattern of
vegetative bud burst in the Trinity River Valley near Salyer, California
[42]:
Elevation Date
< 2,000 feet (610 m) mid-April
2,000 to 3,500 feet (610-1,067 m) mid-May
3,500 to 4,400 feet (1,067-1,341 m) late May
4,500 feet (1,372 m) early June
FIRE ECOLOGY
SPECIES: Lithocarpus densiflora | Tanoak
FIRE ECOLOGY OR ADAPTATIONS :
Tanoak resprouts following fire via dormant buds located on an
underground regenerative organ known as a burl or lignotuber [36,53].
Stored carbohydrates in the burl and an extensive taproot system aid in
a rapid and aggressive postburn recovery [31,53]. Resistance to low
intensity burning is increased in older individuals where the bark may
be from 1 to 3 inches (2.5-7.6 cm) thick [36,42].
POSTFIRE REGENERATION STRATEGY :
survivor species; on-site surviving root crown or caudex
FIRE EFFECTS
SPECIES: Lithocarpus densiflora | Tanoak
IMMEDIATE FIRE EFFECT ON PLANT :
Tanoak is a fire-sensitive species. Aboveground portions are extremely
susceptible to fire mortality [53,54]. The thin bark provides little
insulation from radiant heat which usually kills the cambium around the
base of the stem [30]. As a result, low-intensity ground fires readily
top-kill tanoak seedlings and sapling-sized stems [1,48,49]. Larger,
thicker barked trees occassionally survive light underburning [42].
Bark thickness of mature trees may range from from 1 to 3 inches
(2.5-7.5 cm), sometimes reaching 4 to 5 inches (1-12.5 cm). Bole
injuries usually result following ground fires, however, and vertical
wounds 4 to 10 feet (1.2-3 m) long are common [31,42]. Many older tanoak
trees may initially survive light burns, but bole wounds facilitate the
entry of insects, and disease and most injured trees eventually die
[42].
Long-term survival is most likely in young, vigorous trees where bole
wounds tend to heal over rapidly [42]. In virgin redwood stands in
Redwood National Park, Veirs [54] found the oldest tanoak trees occupying
sites where frequent underburning by indigenous peoples reduced fuel
loadings to the point where only light-intensity ground fires occurred.
Crown fires kill the aerial portions of all tanoak, regardless of age or
size [36,42].
DISCUSSION AND QUALIFICATION OF FIRE EFFECT :
Tanoak is more susceptible to fire mortality when it occurs beneath a
mature conifer overstory. Plants under these conditions are subject to
increased stress and are less able to survive fires than when growing in
a more open environment. Understory tanoak exhibited significantly
lower predawn water potentials than those growing in adjacent open areas
[18]. Late spring (June), high duff consumption underburns
significantly (P<.05) reduced tanoak plant densities in the understory
of mixed conifer stands on sites in the Sierra Nevada [18]. (See fire
case study for more details.)
PLANT RESPONSE TO FIRE :
After fire or cutting, tanoak resprouts from adventitious buds located
on a burl or lignotuber [16,31,42,48]. Most buds are located at or
beneath the ground surface [42]. Unless fires are particularly severe,
nearly all tanoak resprout to some extent during the first postburn
growing season [31].
Tanoak initiates a rapid postburn recovery and is an aggressive
competitor during the early stages of postburn succession [1]. Compared
to the slow growth of suppressed tanoak understories, tanoak sprout
development is dynamic in the postburn environment. Sprout clumps
(aggregation of sprouts originating from a single tanoak individual) are
characterized by large numbers of resprouts and resprouts grow rapidly,
increasing greatly in both height and crown width [14]. If abundant in
the preburn community, tanoak often dominates the initial postburn
vegetation within 3 to 6 years, forming a dense cover which may exceed
49,400 stems/acre (20,000 stems/ha) [49]. Tanoak sprout development on
logged and burned sites in northern California is presented below [42].
# of years after Average sprout Average # of
disturbance height sprouts per clump
(tallest in clump)
ft (m)
2 5.2 (1.6) 27
3 6.8 (2.1) 12
4 7.9 (2.4) 10
5 9.3 (2.8) 10
6 10.3 (3.1) 9
The essentially pure, dense, even-aged sprout stands which frequently
result following fire provide the mutual shading necessary for optimal
tanoak development. Where crowns maintain codominance with one another,
height growth is most rapid and tanoak avoids overtopping by associated
species. Within approximately 20 years of burning, 60 to 90 percent of
the resprouts per sprout clump die as growth is concentrated into
multiple, dominant stems [31]. As succession progresses, preburn stand
density becomes a significant factor influencing both the growth and
continued dominance of tanoak [31].
DISCUSSION AND QUALIFICATION OF PLANT RESPONSE :
Postburn sprouting potential of tanoak is strongly correlated with size
and vigor of the parent tree [42,48,49]. Site quality apparently has
almost no influence on sprout-clump development for at least 6 years
after disturbance [31,48].
Since burl size increases as tanoaks grow, larger stemmed tanoak (> 0.8
inches [2 cm] d.b.h.) usually possess larger burls with increased
numbers of dormant buds [49]. Trees greater than 12 inches (30 cm)
d.b.h. typically support abundant resprouts. Sprout production by
vigorous, large diameter trees is impressive, with as many as 100
resprouts observed on some tanoak individuals. Sprout growth is
intially fueled by portions of the residual root system. Sprout growth
is related to parent tree diameter [31]. On sites in southwestern
Oregon, tanoaks 1 to 4 inches (2 to 10 cm) d.b.h. produced sprout clumps
measuring 4.9 feet (1.5 m) tall and 3.6 feet (1.1 m) wide within 6 years
of logging [48,49]. Trees 8.3 to 11.8 inches (21-30 cm) d.b.h.
produced clumps 7.2 feet (2.2 m) tall with crown widths equalling 8.5
feet (2.6 m). Previous fires and the subsequent entry of pathogens can
substantially reduce the sprouting potential of parent trees greater
than 16 inches (40 cm) d.b.h. [14]. Sprout numbers also tend to be
reduced in older trees where dormant buds are covered by thick bark and
where logging debris is piled against tanoak stumps following
clearcutting [31].
Although tanoak is able to sprout at a very young age, sprouting
potential develops slowly. Younger, smaller stemmed tanoak (< 0.8
inches [2 cm] d.b.h.) sprout much less vigorously than older
individuals. Whereas stems in this smaller size class are usually
destroyed during fire, sprouting potential is related to burl diameter
[49]. Sprouting potential is enhanced as small tanoak age. Older
individuals with larger burls generally produce the most vigorous
sprouts [31]. On sites in southwestern Oregon, small tanoak produced the
following pattern of development within 5 to 6 years after fire or
cutting [48]:
average average average average
age burl sprout number of clump
diameter height sprouts diameter
(mm) inches (cm) inches (cm)
14 - 20 5 - 25 11.2 (28) 4 6.2 (15.6)
28 - 36 26 - 50 25.6 (64) 5 13.9 (34.8)
40 - 52 51 - 75 28.4 (71) 7 14.6 (36.5)
FIRE MANAGEMENT CONSIDERATIONS :
Broadcast burning: Where conifer regeneration is a primary management
concern, broadcast burning is generally an ineffective site preparation
tool following clearcutting in conifer-tanoak stands [30]. Even though
burning delays the recovery of tanoak for approximately one growing
season, removal of logging debris promotes resprouting by exposing basal
buds to solar heating and permits sprouts to grow unimpeded [15,31].
Sites which are particularly prone to the rapid development of a dense
tanoak understory are those where the preburn vegetation consists of low
conifer stocking combined with high tanoak densities [49].
Preharvest underburning: Tanoak sprouting may be more effectively
controlled by preharvest underburning treatments [18,19,49].
Preliminary research indicates that preharvest underburns are effective
in killing at least a portion of the tanoak understory when conducted in
late spring (June) under conditions which result in high duff
consumption [18,19].
Fires aimed at suppressing the tanoak understory can be expected to most
effective when conducted in 30 to 75 year old conifer stands.
Harvesting conifer-tanoak stands over 70 years of age typically results
in a dense cover of tanoak resprouts. On many sites, resprouts from
delayed seedling regeneration are often responsible for high
postdisturbance tanoak coverages [49]. This younger, shrublike
component of the tanoak understory is an abundant (commonly exceeding
3,000 per/ha) but inconspicuous component of many stands prior to
disturbance. Resprouting capabilities are greatly enhanced as
suppressed seedlings increase in age. When released from dense shade,
suppressed tanoak over 70 years of age commonly produce more than 30
sprouts per plant; these typically reach heights of from 3.2 to 6.6 feet
(1 to 2 m) within three years. By comparison, most 40 to 50 year old
tanoak produce an average of only five resprouts ranging from 12 to 27
inches (30 to 68 cm) in height 3 years after burning. Furthermore,
development of a tanoak understory capable of vigorous, postdisturbance
sprouting may take upwards of 100 to 137 years [31]. The sprouting
potential of tanoak develops slowly and substantial tanoak seedling
establishment is often delayed until conifer stands are 20 to 35 years
old. These factors suggest that on sites with 60 to 80 year conifer
rotations, one, well-timed, effective underburn can eliminate tanoak as
a competitor to conifer regeneration for two rotations. Subsequent
tanoak seedling establishment must be controlled in order to realize the
full benefits of burn treatments; resprouted individuals can produce
acorns within at least 9 to 13 years of clearcutting and burning [49].
Hardwood management: Burning should not be utilized as a method of
slash disposal in partially cut hardwood stands where tanoak is managed
for timber production. Instead, logging debris should be lopped and
scattered or piled and burned [30].
FIRE CASE STUDIES
SPECIES: Lithocarpus densiflora | Tanoak
CASE NAME :
Mixed conifer - Sierra Nevada
REFERENCES :
1. Kaufmann, J. B.; Martin, R. E. 1985 [18]
2. Kauffman, J. B.; Martin, R. E. 1985 [19]
SEASON/SEVERITY CLASSIFICATION :
early spring/moderate
late spring/high
early fall/high
late fall/moderate
STUDY LOCATION :
This prescribed burn took place in the Challenge Experimental Forest on
the LaPorte Ranger District, Plumas National Forest. The study site was
located approximately 2.5 miles (4.0 km) southeast of Callenge,
California.
PREFIRE VEGETATIVE COMMUNITY :
Preburn overstory was dominated by ponderosa pine, Douglas-fir, and
sugar pine. Common understory associates included black oak, incense
cedar (Calocedrus decurrens), and bear clover (Chamaebatia foliolosa).
The site is described as productive.
TARGET SPECIES PHENOLOGICAL STATE :
burn phenological state
early spring burns prior to period of active growth
late spring burns coincided with period of active
leaf growth and stem expansion
early fall burns aboveground shrub growth has ceased
late fall burns leaf abscission was occurring in
deciduous shrubs
SITE DESCRIPTION :
Elevation: 3,280 feet (1,000 m)
Aspect: generally west
Slope: 1 to 12 percent
Site index productivity class: I to III
FIRE DESCRIPTION :
season early late early late
fall fall spring spring
duff consumption % 93.6 83.4 91.6 69.7
duff consumption t/ha 111.2 105.8 111.3 72.3
total fuel consumption % 92.1 77.5 82.4 56.2
total fuel consump. t/ha 148.2 117.2 135.8 69.2
duff moisture % 15.7 43.4 30.9 119.5
soil moisture % 11.1 22.3 25.7 44.1
flame length cm 30.5 56.3 97.1 70.9
fireline intensity
(kj m -1 s -1) 21.2 85.9 272.7 125.8
residence time sec. 47.8 51.2 83.6 55.9
FIRE EFFECTS ON TARGET SPECIES :
Density of tanoak (numbers/ha) before and after burning are as follows:
preburn postburn
early spring 2,801 1,400**
late spring 1,934 167**
early fall 1,234 233*
late fall 1,934 600*
* P <0.10
** P <0.05
FIRE MANAGEMENT IMPLICATIONS :
Prescribed, preharvest underburning can be an effective method of
controlling tanoak. Greatest tanoak mortality generally occurs
following high consumption burns irregardless of season of burn.
Increased mortality can be expected when high consumption burns coincide
with seasons of active growth. Late spring (June), high consumption
burns produced highest shrub mortality. This Sierra Nevada site tends
toward the dry end of the moisture range.
REFERENCES
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Index
Related categories for Species: Lithocarpus densiflora
| Tanoak
|
 |
