© 2006 Julie Wakelin
|© 2006 Louis-M. Landry|
In British Columbia, Pacific madrone grows with lodgepole pine (Pinus contorta) . The open woodlands of the San Juan Islands are characterized by Douglas-fir and Pacific madrone in a fescue (Festuca spp.) matrix. Other tree species that may be found on such sites include Rocky Mountain juniper (Juniperus scopulorum), lodgepole pine, and Oregon white oak .
Pacific madrone is a dominant species in the following vegetation types.California:
coast live oak (Q. agrifolia)-Pacific madrone/California hazelnut-blackberry (Corylus cornuta var. californica-Rubus spp.)
interior live oak (Q. wislizenii)-Pacific madrone/poison-oak (Toxicodendron diversilobum)
California black oak (Q. kelloggii)-Pacific madrone-coast live oak 
Pacific madrone-Oregon white oak
Pacific madrone-tanoak 
Douglas-fir-tanoak-Pacific madrone Society of American Foresters cover type 
western hemlock-Douglas-fir-Pacific madrone 
Douglas-fir-Pacific madrone/pink honeysuckle (Lonicera hispidula) 
Douglas-fir-Pacific madrone/American vetch (Vicia americana)
Douglas-fir-Pacific madrone/salal (Gaultheria shallon)
Pacific madrone-lodgepole pine/salal
|© 2004 Charles E. Jones||© 2005 Doreen L. Smith|
Pacific madrone is a broadleaved, sclerophyllous, evergreen tree [1,29,40,88]. Heights range from 16 to 130 feet (5 to 40 m) [13,62,101,138], with diameters up to 2 to 3 feet (0.6-1 m) [7,63]. Single or multiple curved trunks support a broad, spreading crown composed of heavy, irregularly-shaped limbs . The bark is freely exfoliating, peeling off in large, thin scales. Once the outer bark is shed, the remaining bark has a smooth, polished appearance and a distinctive reddish color [13,101,111,138]. Color of young bark varies widely but darkens to a deep red with age; younger stems may range from green to chartreuse, while young trunks are frequently orange. Older portions of the bark become dark, brownish-red in color and are fissured [13,62,101]. The glossy, leathery leaves are arranged alternately on the stem [63,106].
The urn-shaped flowers are borne in showy, terminal clusters [63,91,106]. The fruit is a pea-sized berry consisting of mealy pulp and numerous seeds . At the base of its stem, Pacific madrone has a woody, globe-shaped, underground regenerative organ known as a burl [65,131]. The massive, wide-spreading root system is associated with ericoid mycorrhizae [97,105]. Once established, Pacific madrone is windfirm, drought enduring, and somewhat tolerant of wet, freezing conditions [40,89].RAUNKIAER  LIFE FORM:
McDonald and Tappeiner  describe 3 reproductive modes relative to Pacific madrone: seedlings, seedling-sprouts, and root-crown sprouts. Seedlings originate from seed and their tops have never died back to the ground. Seedling-sprouts also originate from seed, but their tops are less than 2 inches (5.1 cm) in diameter at the ground line, and they have died back and sprouted at least once. The chances of Pacific madrone becoming a seedling-sprout are low, and seedling-sprouts rarely occur in shade environments. Root-crown sprouts originate from burls on top-killed trees more than 2 inches (5.1 cm) in diameter at ground line .
Pollination: Pacific madrone is pollinated by bees [13,18,51]. Hummingbirds have been observed feeding on Pacific madrone blossoms and may also pollinate the flowers .
Breeding system: Pacific madrone has low genetic diversity in British Columbia and is known for multilocus outcrossing .
Seed production: The age at which Pacific madrone seedlings first produce fruit is not recorded in the literature. The minimum seed-bearing age for root crown sprouts is 4 years, but seed production occurs more commonly at 8 years . On the Challenge Experimental Forest, California, initial flower production occurred at age 4 on a "vigorous sprout", resulting in 62 berries. On another sprout clump, the tallest and most vigorous sprout produced 11 flower clusters at age 8 but produced few berries. Seed count ranged from 2 to 37 seeds/berry, with an average of 20 seeds/berry .
A 24-year study estimating seed crops of conifer and hardwood species on a Pacific ponderosa pine site on the Challenge Experimental Forest estimated that the average number of berries on 3 Pacific madrone trees was 49,000/tree, with a range of 13,000 to 108,000/tree during a "very light" seed year. The average number of seeds/berry was 20. Over the 24-year period, Pacific madrone produced 12 seed crops. Two were categorized as "medium-heavy", and 10 were categorized as "very light" .
Seed dispersal: Pacific madrone seeds are dispersed largely by birds, but also by mule deer, rodents, and gravity [13,18,71,82,87].
On the Challenge Experimental Forest, berries are disseminated by a host of consumers, particularly the mourning dove and band-tailed pigeon .
Seed banking: McDonald  states that Pacific madrone has long-term seed dormancy and viability and stays viable for "scores" of years in the soil. When conditions are right (i.e., cool temperatures and adequate moisture), after-ripening is induced and dormancy is broken .
Germination: A cold stratification period is critical for germination of Pacific madrone seeds [58,82,91], because the seeds have strong embryo dormancy . McDonald  identified optimal stratification requirements for Pacific madrone seeds through a series of tests including cold, light, heat, acid, and stratification. Seeds failed to germinate after stratification at freezing temperatures for 24 days, while a 24-day stratification at above-freezing temperatures (36 ± 2 °F (2.2 ± 1.1 °C)) yielded 43% germination. Light was apparently unnecessary for germination of Pacific madrone seeds. Percent sound Pacific madrone seeds that germinated after heat, acid (sulphuric acid), and stratification treatments is provided in the table below. Stratification alone and acid and stratification significantly enhanced germination over those treatments using heat (P=0.05). No stratification caused poor germination. Mold was a constant problem in all treatments and in most cases became worse with longer stratification and germination periods .
|Percent of sound Pacific madrone seeds that germinated after 4 stratification treatments and 5 time periods |
Stratification period (days)
|Stratification||Acid & stratification||Heat & stratification||Heat, acid, & stratification|
In a laboratory study on germination, 2 Pacific madrone populations showed only slight differences in length of time required for stratification. Maleike and Hummel  collected seeds from a high-elevation and a sea-level source. The seeds were stratified at 39 °F (4 °C) for 0, 20, 40, 60, and 80 days. Percent germination increased with increasing time in cold stratification up to 60 days. After 60 days there was a decline in percent germination with both seed sources. Maximum germination for the sea-level seeds was reached at both 40 and 60 days. The seeds from the high-elevation seed source reached highest germination at 60 days .
Germination of seeds not separated from the berry was found, in a laboratory study, to be poor and intermittent. Berries were stratified in a refrigerator for 45 days and underwent subsequent germination tests. Seedlings did not readily disengage from the berry and seed coat, and there was heavy mortality from fungi. In field trials on the Challenge Experimental Forest, if the berries and seed survived long enough to germinate (i.e., not eaten by birds, rodents, etc.), many seedlings were killed by damping-off and root-rotting fungi . Fungi appear to be a major problem in natural and artificial regeneration of Pacific madrone.
Seed germination is discouraged by low light intensities under a closed canopy; therefore, Pacific madrone may not reproduce satisfactorily under dense forest conditions .
Seedling establishment/growth: Disturbance favors seedling establishment of Pacific madrone [82,92,132]. Survival rates of artificial Pacific madrone regeneration were observed on 3 types of Douglas-fir-ponderosa pine stands in the Siskiyou Mountains of southwestern Oregon. The 3 stands were differentiated as: clearcut, 5 to 14 years old; a young conifer-hardwood stand, 50 to 80 years old; and an old conifer-hardwood stand, 150 to 220+ years old. Seeds were sown in December at each location. One lot was sown on bare mineral soil protected by a cage, and 1 lot each on unprotected plots on undisturbed forest floor and bare mineral soil. Germinants began to emerge in early March, with more than 90% of the seedlings appearing within 1 month. Fewer seedlings emerged on unprotected plots than on protected plots due to predation of seed. Seedlings began to die immediately after emergence. Average survival at the end of the 1st summer was significantly lower (P=0.05) in old stands (5%-14%) and young stands (8%-12%) than in clearcuts (32%-34%). On most plots all seedlings had died within 1 year, with 1st-year mortality ranging between 90% and 100%. Causes of seedling mortality, in order of importance, were drought, litterfall covering small seedlings during fall months, damping-off fungi, invertebrate browsers (mainly slugs) in both young and old stands, and spring and fall frost, common in the clearcuts. At the end of the 2nd year, survival ranged from less than 1% to 3% in the young and old stands and after 2 and 3 years in the clearcuts, survival ranged from 5% to 12%. In this study, success of seedlings was dependent on disturbance to the forest floor and reduced litterfall, as indicated by the higher survival in clearcut stands . McDonald  stated that the bare mineral soil created by some silvicultural methods is conducive to seedling survival and noted little natural regeneration of Pacific madrone in an undisturbed pure hardwood stand.
Most Pacific madrone seedlings are found in partial shade on bare mineral soil . On recently logged redwood stands in northern California, Pacific madrone established in open environments on relatively hot, dry sites with thin, rocky soil . Seedling establishment is minor in stands with low light, heavy litterfall, damping-off fungi, and browsing invertebrates on the forest floor, all of which kill new seedlings [82,92,105,133,142]. High soil and air temperatures and frost heaving also kill Pacific madrone germinants on exposed microsites in clearcuts. Many Pacific madrone seedlings begin development in heavy organic litter in shade. The heavy organic layer inhibits the moisture-seeking root from penetrating to mineral soil, causing high mortality from fungi and drought .
Early growth of Pacific madrone seedlings is slow. In the Santa Cruz Mountains, California, length of 6-month-old seedlings growing in the sun was 1.6 inches (4 cm) for shoots and 4 inches (10 cm) for roots. Seedlings growing in a shady environment had shoots that were 1 inch (3 cm) and roots measuring 1.6 inches (4 cm). Two-year-old seedlings in the Sierra Nevada averaged 3.5 inches (9 cm) tall [82,92].
Vegetative regeneration: Pacific madrone sprouts from the burl after damage by cutting, fire, or disease [36,59,66,89,131]. It is unknown how early the burl develops on seedlings .
In its southern range in southwestern Oregon and California, Pacific madrone is often associated with dry foothills, wooded slopes and canyons [101,122]. In California, elevations range from 300 to 4,000 feet (91-1,220 m) . Pacific madrone is common above 3,900 feet (1,200 m) in the San Lucia Range of central California . A common component of coastal redwood and mixed-evergreen forests, Pacific madrone reaches greatest stature and abundance on dry sites at low to moderate elevations along the east slope of the Coast Ranges and in the Siskiyou Mountains [48,108,124]. At the southernmost end of its range in the Transverse and Peninsular ranges, Pacific madrone is found from 2,000 to 3,500 feet (610-1,100 m) elevation .
In its northern range, Pacific madrone grows at or near sea level and inhabits mountain slopes up to 3,000 feet (915 m) . Increased regional rainfall apparently allows Pacific madrone to occupy drier habitats than in mixed-evergreen forests . Greatest abundance is usually attained on sites unfavorable to conifer growth [13,42]. Pacific madrone is widespread west of the Cascade Range in Oregon and Washington and is associated with relatively hot, dry lowland sites within coast Douglas-fir and western hemlock forests [43,148]. Pacific madrone communities on Sucia Island, Washington, are located on south-facing ridges where winds are moderate, temperatures somewhat high, and soil moisture low. These sites are protected from extreme wind by windward ridges, but abundant solar radiation strikes the slopes . On the Willamette, Mt Hood, and Siuslaw National Forests of western Oregon, Pacific madrone inhabits dry sites on ridgetops and south-facing slopes up to 5,000 feet (1,500 m) in elevation . Towards the northern edge of its distribution in southern British Columbia and northwestern Washington, Pacific madrone is generally restricted to areas along the immediate coast . The only broadleaved evergreen tree native to Canada [63,74], Pacific madrone rarely extends inland more than 5 miles (8 km) in southern British Columbia [43,63,74]. Sites consist of rocky bluffs along the seacoast; elevations do not exceed 1,000 feet (300 m) [54,63].
Soils: Pacific madrone grows on a variety of soil types, and tree health varies with soil type . Pacific madrone is most abundant on rocky sites, such as bluffs, that are "somewhat excessively" drained [2,13]. Soils supporting Pacific madrone usually exhibit low moisture content throughout most of the summer. Pacific madrone grows on glacial tills or shallow rocky soils in the northern portion of its range. Soils may also be fine textured, ranging from loam to clay loam. Towards the southern end of its distribution, soils are often derived from granite, quartz diorite, sandstone, or shale .
Climate: Pacific madrone is restricted to areas having mild oceanic winters; however, temperature and moisture regimes vary considerably throughout its range. Annual precipitation may range from 15 to 166 inches (380-4,220 mm), mostly as rain. Temperature extremes are from -6 to 115 °F (-21 to 46 °>C) [13,40,92,135].
Pacific madrone is drought tolerant  and has low tolerance to frost. It can be damaged or even killed if it endures long periods of frost or severe frost (<14 °F (-10 °C)) .
Pacific madrone has moderate to low shade tolerance [13,16,64,74] and is considered an early-successional hardwood after timber harvest, fire [9,19,55], and other disturbances. Pacific madrone does not generally establish in shade and is usually absent from the understory of mixed-evergreen forests. A study of a Douglas-fir-tanoak forest in Mendocino County, California, revealed no Pacific madrone recruitment in the understory. Most shaded Pacific madrone died .
The level of shade tolerance can vary depending on Pacific madrone's north-south range. In the southern portion of its range, Pacific madrone seedlings need partial shade for establishment. As Pacific madrone trees age, the need for light increases, and older trees require top light for survival. In British Columbia Pacific madrone has low shade tolerance [74,92], so Douglas-fir dominates over Pacific madrone in climax stages .
Pacific madrone is likely more often subclimax than climax in successional status . Pacific madrone can be eliminated during the stem exclusion stage of succession, but it is possible for it to survive stem exclusion and persist into the old-growth stage . Pacific madrone has been documented in climax forests [15,121] and is classified as a "major climax species" in the western hemlock-Douglas-fir-Pacific madrone association on the Gifford Pinchot National Forest, Washington .
Pacific madrone is considered a fire-dependent, seral species in redwood stands of northern coastal California .
Pacific madrone typically bears flowers in May, but may flower in March and April at low elevations [13,82,92]. It flowers from April to May on the Willamette, Mt Hood, and Siuslaw National Forests of western Oregon . In June, the second-year leaves turn orange to red and begin to fall shortly after the new crop of leaves has fully grown. Bark is shed all summer. Berry clusters ripen in autumn and persist into December . On the Challenge Experimental Forest, Pacific madrone berries mature from mid-September to mid-October . The table below gives generalized seasonal development of southern and northern populations of Pacific madrone.
|Generalized trends in the phenological development of Pacific madrone |
|Southern range||Northern range|
|Leaf bud swelling begins||February||late March|
|Flower bud swelling begins||March||May|
|Second-year leaves fall||June||June-July|
Postfire regenerative adaptations include establishment from prolific sprouts and from seed . Following fires that kill aerial stems, Pacific madrone sprouts from dormant buds on the burl [40,59,94,140]. The burl also serves as a source of stored carbohydrates for the sprouts, which rapidly occupy the initial postfire environment [65,86]. Repeated top-kill by fire encourages burl development, enhancing Pacific madrone survival [14,70].
Exposed mineral soil seedbeds and light canopy densities associated with recent burns are conducive to Pacific madrone seedling establishment [14,105,132].
Fire regime: Forests where Pacific madrone occurs were historically characterized by both understory and mixed-severity fires prior to fire exclusion. Oak-Pacific madrone-Douglas-fir and redwood forests, where Pacific madrone occurs, historically experienced understory fires at intervals between 5 and 25 years. Historic fires on some sites were caused mainly by Native American burning [11,12,78]. Remote, steep areas of the redwood type were also likely associated with a mixed fire regime . A redwood-Douglas-fir stand in northern coastal California had fires approximately every 50 years over the past 250 years . Fire typically burned through Douglas-fir-tanoak forests in northern California at 5- to 50-year intervals, killing small saplings and occasional canopy trees. These forests now have very infrequent fires . Douglas-fir-tanoak-Pacific madrone and Douglas-fir-hardwood cover types were characterized by a mixed-severity fire regime, with the former having less than 35-year fire-return intervals . In Douglas-fir/hardwood forests of the Pacific Northwest, the severity of fires varied widely, with many burning at low to moderate severity prior to settlement .
Stuart and Stephens  review fire regime characteristics for Douglas-fir-tanoak forests that Pacific madrone commonly occurs in. Presettlement fire-return intervals averaged from 10 to 16 years due to the warm, dry climate of inland locations and increased lightning activity at high elevations. In the North Coast Ranges, the primary ignition source was Native Americans. There is little information available on the size and severity of fires in Douglas-fir-tanoak prior to settlement. Areas subject to Native American burning experienced low fire severity. In other areas, fire severity varied spatially and temporally across the landscape, resulting in a complex mosaic of mostly multiaged stands of varying sizes. Fires in interior sites spread more extensively than those closer to redwood forests. Surface fires were common and were intermixed with areas that supported passive and/or active crown fires. In the Six Rivers National Forest, California, surface fires are a normal occurrence in all-aged, all-sized old-growth Douglas-fir-tanoak forests. In old-growth Douglas-fir-tanoak forests, the density of understory trees and shrubs has increased since presettlement times, creating greater vertical fuel continuity and increasing the likelihood that a surface fire could burn into the crowns. In young stands that have been logged or experienced stand-altering wildfire, fire-return intervals are now longer, and greater fire severity is possible because of increased fuel loading .
Pacific madrone is common in redwood forests. Fire in redwood forests typically burned in the summer and early fall with variable fire-return intervals. Wetter sites in the northern portion of redwood's range had longer fire-return intervals, ranging from 125 to 500 years. Drier sites in southern locales experienced shorter fire-return intervals, between 6 and 44 years. Some stands had intervals of 1 to 2 years due to regular burning by Native Americans. From 1950 to 2003, fire-return intervals for the northern, central, and southern redwood forests for fires larger than 330 acres (134 ha) were 1,083, 717, and 551 years, respectively. On average, redwood forests experienced moderate-severity surface fires that consumed irregular patches of surface fuel and understory vegetation. Occasional passive crown fires occurred at the southern and eastern edges of its range. Throughout the north-south range of redwood forests, mean fire severities were lowest in the coolest, wettest regions and highest in the warmer, drier areas. The current increase of available fuel and increasing horizontal and vertical fuel continuity may increase the chances for higher severity fires (review by Stuart and Stephens ).
White fir (Abies concolor) forests in the Coast Ranges of northwestern California, in which Pacific madrone can occur, had a presettlement average fire-return interval of 27 years, with a range of 12 to 161 years. The average fire-return interval has increased to 74 years since the exclusion of fire .
Fire scar, tree age, and basal area distributions were used to assess fire history in 3 Douglas-fir/hardwood stands in the Klamath National Forest, California (see table below). Fire-return intervals changed little from the presettlement era to the settlement era but increased in the fire exclusion era. The upper canopy of was dominated by Douglas-fir with scattered stems of sugar pine (Pinus lambertiana). The lower canopy was dominated by tanoak, Pacific madrone, and canyon live oak .
|Means and ranges (in years) of fire intervals for 3 Douglas-fir/hardwood sites in the Klamath National Forest, California, for 4 different time periods |
|Site 1 (n = 11)|
|Site 2 (n = 19)|
|Site 3 (n = 13)|
The following table provides fire regime information on vegetation communities in which Pacific madrone may occur. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".
|Fire regime information on vegetation communities in which Pacific madrone may occur. For each community, fire regime characteristics are taken from the LANDFIRE Rapid Assessment Vegetation Models . These vegetation models were developed by local experts using available literature, local data, and/or expert opinion as documented in the PDF file linked from each Potential Natural Vegetation Group listed below. Cells are blank where information is not available in the Rapid Assessment Vegetation Model.|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval
|Oregon white oak-ponderosa pine||Replacement||16%||125||100||300|
|Surface or low||81%||25||5||30|
|Oregon white oak||Replacement||3%||275|
|Surface or low||78%||12.5|
|Sitka spruce-western hemlock||Replacement||100%||700||300||>1,000|
|Douglas-fir (Willamette Valley foothills)||Replacement||18%||150||100||400|
|Surface or low||53%||50||20||80|
|Oregon coastal tanoak||Replacement||10%||250|
|Douglas-fir-western hemlock (dry mesic)||Replacement||25%||300||250||500|
|Douglas-fir-western hemlock (wet mesic)||Replacement||71%||400|
|Mixed conifer (southwestern Oregon)||Replacement||4%||400|
|Surface or low||67%||22|
|California mixed evergreen (northern California)||Replacement||6%||150||100||200|
|Surface or low||64%||15||5||30|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval
|California oak woodlands||Replacement||8%||120|
|Surface or low||91%||10|
|California mixed evergreen||Replacement||10%||140||65||700|
|Surface or low||32%||45||7|
|Surface or low||98%||20|
|Mixed conifer (North Slopes)||Replacement||5%||250|
|Surface or low||88%||15||10||40|
|Mixed conifer (South Slopes)||Replacement||4%||200|
|Surface or low||80%||10|
|Mixed evergreen-bigcone Douglas-fir (southern coastal)||Replacement||29%||250|
Replacement=Any fire that causes greater than 75% top removal of a vegetation-fuel type, resulting in general replacement of existing vegetation; may or may not cause a lethal effect on the plants.
Mixed=Any fire burning more than 5% of an area that does not qualify as a replacement, surface, or low-severity fire; includes mosaic and other fires that are intermediate in effects
Surface or low=Any fire that causes less than 25% upper layer replacement and/or removal in a vegetation-fuel class but burns 5% or more of the area. [57,76].
|Photo taken by John Smiley, 2000|
IMMEDIATE FIRE EFFECT ON PLANT:
Fire top-kills most Pacific madrone [4,6,25] of all sizes, but they generally only die back to the burl [14,92]. Some large Pacific madrones may survive moderately-severe fire but sustain bole damage that leaves fire scars [25,31].
DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
Aboveground portions of Pacific madrone are very susceptible to fire damage [11,14,32,94,143]. Thin bark provides little insulation from radiant heat, which usually kills the cambium around the base of the stem . Even the thicker bark at the base of old trees shields them little ; however, it may explain how some Pacific madrones survive with only moderate damage after low-severity fire. Individuals that withstand fire have moderate susceptibility to secondary attack by insects or disease , which may result in mortality.
PLANT RESPONSE TO FIRE:
Sprout growth from top-killed trees is the primary mode of reproduction following fire . Following fires that kill aerial stems, Pacific madrone initiates rapid postfire recovery by sprouting from adventitious buds located on the burl [1,10,21,25,44,65,90,131]. Seedlings on the forest floor are either killed or put forth a few sprouts from a rudimentary burl. Many seedlings die because the growth rate of sprouts from top-killed trees is much greater, and the sprouts overtop the seedlings.
Fire favors Pacific madrone seedling establishment. Mineral soil provides a favorable seedbed, and lower canopy densities of the initial postfire environment are conducive to the successful establishment and growth of seedlings [105,132]. Availability of seed from crowns depends on fire severity . Any seeds stored in the soil seed bank are likely killed due to their sensitivity to heat . Off-site seed is also dispersed into burns by mammals and birds .DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
|Pacific madrone sprout development after summer wildfire in northwestern California |
|Height of tallest sprout in clump (feet)||Crown diameter of sprout clump (feet)||Sprouts/clump|
|Time since burning (August 1951)||Average*||Range||Average||Range||Average||Range|
|1 year (November 1952)||4.7||1.6-7.6||4.5||0.8-8.9||17||1-47|
|2 years (October 1953)||7.7||3.2-11.5||6.8||2.0-13.7||16||1-47|
|3 years (September 1954)||10.1||4.9-14.8||7.6||2.8-16.5||13||1-32|
|*n=50 except for 3rd year where n=48|
A recently burned site at Fort Lewis, Washington, had abundant Pacific madrone seedlings in addition to Pacific madrone sprouts .
The Research Project Summary of Kauffman and Martin's [68,69] study provides information on prescribed fire and postfire response of many species in mixed-conifer forests, including Pacific madrone. Pacific madrone occurred on the Challenge Experimental Forest, which was 1 of 3 study sites. Sprouts and/or seedlings were observed in postfire year 1 on two fall burns but were not found in postfire year 2 [68,69].FIRE MANAGEMENT CONSIDERATIONS:
Prescribed burning: Pacific madrone seedlings establish readily following logging and burning of conifer-hardwood stands . Low-severity underburning may minimize Pacific madrone seedling establishment, thereby reducing the density of Pacific madrones capable of sprouting after future disturbances.
Control: McDonald and others  suggest that burning should not be used as a method of slash disposal in partially cut hardwood stands where Pacific madrone is managed for timber production. Instead, they recommend that logging debris be either lopped and scattered or piled .Wildlife management: Burning initially increases the palatability of Pacific madrone browse; sprouts are utilized for up to 2 growing seasons after fire [33,140].
Pacific madrone typically grows with mixtures of evergreen and deciduous hardwood species. Mixed stands are highly diverse in structure and composition and provide habitat for numerous wildlife species [89,108].
Palatability/nutritional value: Palatability of Pacific madrone foliage ranks from low to moderately high, depending on conditions. The mature leaves are almost always neglected by browsing animals, whereas the young leafy sprouts are eaten by big game, domestic sheep and goats, deer, and occasionally cattle, when there is a shortage of more palatable vegetation [33,53,113,122]. Pacific madrone leaves provide forage for the dusky-footed woodrat . Cattle and deer browse the seedlings . In California, the Columbian black-tailed and California mule deer browse twigs and foliage [81,116]. Pacific madrone is given a browse rating of fair to useless for mule deer, poor to useless for cattle, domestic sheep and goats, and useless to horses in California .
In British Columbia, Pacific madrone is a high-importance winter forage plant for Sitka black-tailed deer, of moderate importance for Roosevelt elk, and of low importance for white-tailed deer, mountain goats, bighorn sheep, Rocky Mountain elk, moose, and caribou . Leaves are eaten by Columbian black-tailed deer on Vancouver Island, British Columbia; on rock-bluff communities where Pacific madrone is abundant, it is a major food species during the winter .
Pacific madrone berries are an important food for deer, birds, and other small mammals because they are produced in large quantities and may persist on the tree in winter, when alternative food sources are limited [33,58,122]. The berries are an important food for the dark-eyed junco, fox sparrow, varied thrush, band-tailed pigeon, quail, and long-tailed chat [13,52,81,125].
Cover value: Mixed stands of hardwoods and conifers in which Pacific madrone occurs provide thermal, hiding, and escape cover for big game and small mammals, and perching sites for a variety of bird species [24,89,118]. Both open-nesting and cavity-nesting birds utilize Pacific madrone. Preliminary research on cavity-nesting species within mixed-evergreen forests in northwestern California indicates that Pacific madrone is selected as a nest tree at a higher rate than its availability would suggest. Trees greater than 12 inches (30 cm) in DBH are an important habitat component for primary cavity-nesting species such as the red-breasted sapsucker and hairy woodpecker . Secondary cavity nesters such as the acorn woodpecker, downy woodpecker, mountain chickadee, house wren, and western bluebird also use Pacific madrone.VALUE FOR REHABILITATION OF DISTURBED SITES:
Historically, West Coast tribes ate Pacific madrone berries and fashioned eating utensils from the bulbous roots [13,56]. The leaves have been reported to possess medicinal properties . Fruit of Pacific madrone can be eaten raw, boiled, or steamed. Berries can be stored for a long time if boiled and dried .
OTHER MANAGEMENT CONSIDERATIONS:
Pacific madrone can reduce conifer growth on previously logged or burned sites and is often considered an undesired competitor [47,48,70,95,107]. The sprouts can reach up to 12 feet (3.7 m) in 3 growing seasons and are capable of creating dense brushfields, hindering conifer establishment [47,48,70]. Control of Pacific madrone is often needed to promote growth of more "valuable" trees such as Douglas-fir . Herbicide applications are a commonly used method to set back Pacific madrone sprouting. Young sprouts are susceptible 2,4-D [28,47,48,70]. Cut-surface applications of herbicides gave acceptable control for 10 years following application. Ten years after treatment, the Douglas-fir site not receiving herbicide control had a mean basal area of 7.2 cm². Basal growth of Douglas-fir receiving the benefit of overstory control increased between 260% and 451%, depending on the herbicide used . The number of postharvest sprouts of Pacific madrone can be reduced by choosing what season cutting is done. More sprouts appeared after April cutting than February or July cutting. At times, Pacific madrone control may be needed to increase forage production .
In the past, leaching from Pacific madrone litter was thought to be allelopathic. In a study by Rose and others , root growth of Douglas-fir seedlings was not inhibited by Pacific madrone litter . Excellent natural regeneration of Douglas-fir often occurs under Pacific madrone canopies, as noted by Minore , but the effects of Pacific madrone duff on Douglas-fir regeneration are not clear. There were no significant differences in conifer regeneration, growth, or cover of associated species among seedbeds of Pacific madrone duff, conifer duff, or mineral soil during 10 growing seasons. If Pacific madrone litter does affect Douglas-fir regeneration, it is because of other influences , possibly the reduction in mycorrhizal tips on Douglas-fir seedling roots .
Disease: Pacific madrone has low resistance to disease and is host to many pathogens that may lead to tree mortality. Pacific madrone can suffer from foliar diseases caused by a variety of fungal species and is susceptible to heart rot, butt rot, and stem cankers . A fungal leaf blister disease caused by Exobasidium vacinii occurs on Pacific madrone leaves. This disease is not thought to significantly reduce tree growth, but it does reduce the aesthetic value of the tree. Phytophthora cactorum is a lethal canker disease of Pacific madrone that results in root and butt rots [23,136].
All ages and sizes of Pacific madrone are susceptible to dieback and mortality from Arbutus canker, a disease caused by the fungus Nattrassia mangiferae. The fungus infects the phloem and vascular cambium and causes shoot blight. Greater weakening of the trees through defoliation is caused by a secondary opportunistic pathogen, Fusicoccum aesculi, which causes dieback and gives the limbs and twigs a burned appearance. The branches and terminal buds that are killed by fungi are unable to produce more foliage. The tree does sprout from dormant buds on the burl and grows new shoots, which are often killed by N. mangiferae .
Pacific madrone is affected by Sudden Oak Death (Phytophthora ramorum). Sudden Oak Death causes a variety of foliar and branch symptoms, significant dieback, and mortality [41,45,46,102,114,115].
The madrone canker (Botryosphaeria dothidea) greatly reduces seed production and causes dieback and death of Pacific madrone [91,92]. Annosus root rot can cause mortality to Pacific madrone . For an extensive list on the fungal pathogens that effect Pacific madrone, see Elliott .
Many Pacific madrones were sampled around the Seattle/Puget Sound area to gauge the effect of urban development and disturbance and whether they facilitated disease transmission and tree demise. Thinning stands, soil loss and compaction, and a host of urban impacts increased susceptibility to disease. Dense stands of Pacific madrone were less infected, and it was predicted that an increase in the proportion of seriously diseased trees would occur if forest stands were broken up .
Other Threats: Scotch broom (Cytisus scoparius) and gorse (Ulex europaeus) are invasive, nonnative plant species that compete with native forest vegetation for space, nutrients, and water. They are a threat to the sustainability of Canada's rarest forest ecosystem, the Oregon white oak-Pacific madrone ecosystem on southeastern Vancouver Island and the southern Gulf Islands .
1. Ackerly, David. 2004. Functional strategies of chaparral shrubs in relation to seasonal water deficit and disturbance. Ecological Monographs. 74(1): 25-44. 
2. Adams, A. B. 1999. Arbutus menziesii and soils of the Puget Trough, Washington. In: Adams, A. B.; Hamilton, Clement W., eds. The decline of the Pacific madrone (Arbutus menziesii Pursh): Current theory and research directions: Proceedings of the symposium; 1995 April 28; Seattle, WA. Seattle, WA: Save Magnolia's Madrones, Center for Urban Horticulture, Ecosystems Database Development and Research: 140-146. 
3. Adams, A. B.; Harvey, F. J.; Crooks, W. T.; Williston, P.; Cholvin, V.; Wilson, R. F. 1999. Habitat physical structure and Arbutus menziesii status in Seattle, Washington. In: Adams, A. B.; Hamilton, Clement W., eds. The decline of the Pacific madrone (Arbutus menziesii Pursh): Current theory and research directions: Proceedings of the symposium; 1995 April 28; Seattle, WA. Seattle, WA: Save Magnolia's Madrones, Center for Urban Horticulture, Ecosystems Database Development and Research: 50-82. 
4. Agee, James K. 1991. Fire history of Douglas-fir forests in the Pacific Northwest. In: Ruggiero, Leonard F.; Aubry, Keith B.; Carey, Andrew B.; Huff, Mark H., technical coordinators. Wildlife and vegetation of unmanaged Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-285. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station: 25-33. 
5. Agee, James K. 1993. Fire ecology of Pacific Northwest forests. Washington, DC: Island Press. 493 p. 
6. Agee, James K.; Edmonds, Robert L. 1992. Appendix E: Forest protection in the Pacific Northwest. In: U.S. Department of Interior, Recovery Plan for the northern spotted owl. Seattle, WA: University of Washington, College of Forest Resources: 56 p. 
7. Alden, Harry A. 1995. Hardwoods of North America, [Online]. In: Gen. Tech. Rep. FPL-GTR-83. Madison, WI: U.S. Department of Agriculture, Forest Service, Forest Products Laboratory (Producer). 136 p. Available: http://www.fpl.fs.fed.us/documnts/fplgtr/fplgtr83.pdf [2004, January 6]. 
8. Allen, Barbara H.; Holzman, Barbara A.; Evett, Rand R. 1991. A classification system for California's hardwood rangelands. Hilgardia. 59(2): 1-45. 
9. Amaranthus, M. P.; Li, C. Y.; Perry, D. A. 1990. Influence of vegetation type and madrone soil inoculum on associative nitrogen fixation in Douglas-fir rhizospheres. Canadian Journal of Forest Research. 20: 368-371. 
10. Ammirati, Joseph Frank, Jr. 1967. The occurrence of annual and perennial plants on chaparral burns. San Francisco, CA: San Francisco State College. 140 p. Thesis. 
11. Arno, Stephen F. 2000. Fire in western forest ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 97-120. 
12. Arno, Stephen F.; Allison-Bunnell, Steven. 2002. Flames in our forest: disaster or renewal? Washington, DC: Island Press. 227 p. 
13. Arno, Stephen F.; Hammerly, Ramona P. 1977. Northwest trees. Seattle, WA: The Mountaineers. 222 p. 
14. Atzet, Thomas; Wheeler, David L. 1982. Historical and ecological perspectives on fire activity in the Klamath Geological Province of the Rogue River and Siskiyou National Forests. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 16 p. 
15. Bailey, Arthur Wesley. 1966. Forest associations and secondary succession in the southern Oregon Coast Range. Corvallis, OR: Oregon State University. 166 p. Thesis. 
16. Baker, Frederick S. 1949. A revised tolerance table. Journal of Forestry. 47: 179-181. 
17. Balfour, Patty M. 1989. Effects of forest herbicides on some important wildlife forage species. Victoria, BC: British Columbia Ministry of Forests, Research Branch. 58 p. 
18. Beland, J. D.; Krakowski, J.; Ritland, C. E.; Ritland, K.; El-Kassaby, Y. A. 2005. Genetic structure and mating system of northern Arbutus menziesii (Ericaceae) populations. Canadian Journal of Botany. 83(12): 1581-1589. 
19. Borchers, Susan L.; Perry, David A. 1990. Growth and ectomycorrhiza formation of Douglas-fir seedlings grown in soils collected at different distances from pioneering hardwoods in southwest Oregon clear-cuts. Canadian Journal of Forest Research. 20(6): 712-721. 
20. Borchert, Mark. 1994. SRM 202: Coast live oak woodland. In: Shiflet, Thomas N., ed. Rangeland cover types of the United States. Denver, CO: Society for Range Management: 12-13. 
21. Brandegee, T. S. 1891. The vegetation of "burns". Zoe. 2: 118-122. 
22. Bullen, Susan; Wood, R. E. 1979. Fomes annosus on Pacific madrone. Plant Disease Reporter. 63(10): 844. 
23. Byther, Ralph S. 1999. Some observations of madrone diseases. In: Adams, A. B.; Hamilton, Clement W., eds. The decline of the Pacific madrone (Arbutus menziesii Pursh): Current theory and research directions: Proceedings of the symposium; 1995 April 28; Seattle, WA. Seattle, WA: Save Magnolia's Madrones, Center for Urban Horticulture, Ecosystems Database Development and Research: 34-37. 
24. Carey, Andrew B. 1991. The biology of arboreal rodents in Douglas-fir forests. Gen. Tech. Rep. PNW-276. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 46 p. 
25. Chappell, Christopher B.; Giglio, David F. 1999. Pacific madrone forests of the Puget Trough, Washington. In: Adams, A. B.; Hamilton, Clement W., eds. The decline of the Pacific madrone (Arbutus menziesii Pursh): Current theory and research directions: Proceedings of the symposium; 1995 April 28; Seattle, WA. Seattle, WA: Save Magnolia's Madrones, Center for Urban Horticulture, Ecosystems Database Development and Research: 2-11. 
26. Cole, David. 1977. Ecosystem dynamics in the coniferous forest of the Willamette Valley, Oregon, U.S.A. Journal of Biogeography. 4: 181-192. 
27. Colwell, Jr., Wilmer L. 1980. Knobcone pine. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 124-125. 
28. Conard, Susan G.; Emmingham, W. H. 1984. Herbicides for forest brush control in southwestern Oregon. Corvallis, OR: Oregon State University, College of Forestry. 7 p. 
29. Cooper, William Skinner. 1922. The broad-sclerophyll vegetation of California: An ecological study of the chaparral and its related communities. Publ. No. 319. Washington, DC: The Carnegie Institution of Washington. 145 p. 
30. Cowan, Ian McTaggart. 1945. The ecological relationships of the food of the Columbian black-tailed deer, Odocoileus hemionus columbianus (Richardson), in the coast forest region of southern Vancouver Island, British Columbia. Ecological Monographs. 15(2): 110-139. 
31. Dale, Virginia H.; Hemstrom, Miles A.; Franklin, Jerry F. 1984. The effect of disturbance frequency on forest succession in the Pacific Northwest. In: New forests for a changing world: Proceedings of the 1983 convention of The Society of American Foresters; 1983 October 16-20; Portland, OR. Bethesda, MD: Society of American Foresters: 300-304. 
32. Dale, Virginia H.; Hemstrom, Miles; Franklin, Jerry. 1986. Modeling the long-term effects of disturbances on forest succession, Olympic Peninsula, Washington. Canadian Journal of Forest Research. 16: 56-57. 
33. Dayton, William A. 1931. Important western browse plants. Misc. Publ. No. 101. Washington, DC: U.S. Department of Agriculture. 214 p. 
34. del Moral, Roger; Cates, Rex G. 1971. Allelopathic potential of the dominant vegetation of western Washington. Ecology. 52(6): 1030-1037. 
35. Elliott, Marianne. 1999. Diseases of Pacific madrone. In: Adams, A. B.; Hamilton, Clement W., eds. The decline of the Pacific madrone (Arbutus menziesii Pursh): Current theory and research directions: Proceedings of the symposium; 1995 April 28; Seattle, WA. Seattle, WA: Save Magnolia's Madrones, Center for Urban Horticulture, Ecosystems Database Development and Research: 42-49. 
36. Elliott, Marianne; Edmonds, Robert L.; Mayer, Scott. 2002. Role of fungal diseases in decline of Pacific madrone. Northwest Science. 76(4): 293-303. 
37. Fiddler, Gary O.; McDonald, Philip M. 1984. Alternatives to herbicides in vegetation management: a study. In: Proceedings of the 5th forest vegetation management conference; [Date of conference unknown]; Sacramento, CA. Redding, CA: The Conference: 115-126. 
38. Finch, Sherman J.; McCleery, Dick. 1980. California coast live oak. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 127-128. 
39. Fonda, R. W.; Bernardi, J. A. 1976. Vegetation of Sucia Island in Puget Sound, Washington. Bulletin of the Torrey Botanical Club. 103(3): 99-109. 
40. Fowells, H. A., compiler. 1965. Silvics of forest trees of the United States. Agric. Handb. 271. Washington, DC: U.S. Department of Agriculture, Forest Service. 762 p. 
41. Frankel, Susan. 2002. Sudden oak death caused by a new species, Phytophthora ramorum, [Online]. In: Pest Alert: NA-PR-06-01. St. Paul, MN: U.S. Department of Agriculture, Forest Service, State and Private Forestry, Northeastern Area (Producer). 3 p. Available: http://www.na.fs.fed.us/spfo/pubs/pest_al/sodwest/sodwest.htm [2004, April 27]. 
42. Franklin, Jerry F. 1979. Vegetation of the Douglas-fir region. In: Heilman, Paul E.; Anderson, Harry W.; Baumgartner, David M., eds. Forest soils of the Douglas-fir region. Pullman, WA: Washington State University, Cooperative Extension Service: 93-112. 
43. Franklin, Jerry F.; Dyrness, C. T. 1973. Natural vegetation of Oregon and Washington. Gen. Tech. Rep. PNW-8. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 417 p. 
44. Franklin, Jerry F.; Swanson, Frederick J.; Harmon, Mark E.; Perry, David A.; Spies, Thomas A.; Dale, Virginia H.; McKee, Arthur; Ferrell, William K.; Means, Joseph E.; Gregory, Stanley V.; Lattin, John D.; Schowalter, Timothy D.; Larsen, David. 1991. Effects of global climatic change on forests in northwestern North America. Northwest Environmental Journal. 7(2): 233-254. 
45. Garbelotto, Matteo. 2004. Sudden oak death: a tale of two continents. Outlooks on Pest Management. April: 85-89. 
46. Garbelotto, Matteo; Davidson, Jennifer M.; Ivors, Kelly; Maloney, Patricia E.; Huberli, Daniel; Koike, Steven T.; Rizzo, David M. 2003. Non-oak native plants are main hosts for sudden oak death pathogen in California. California Agriculture. 57(1): 18-23. 
47. Gratkowski, H. 1975. Silvicultural use of herbicides in Pacific Northwest forests. Gen. Tech. Rep. PNW-37. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 44 p. 
48. Gratkowski, H. 1978. Herbicides for shrub and weed control in western Oregon. Gen. Tech. Rep. PNW-77. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 48 p. 
49. Griffin, James R. 1975. Plants of the highest Santa Lucia and Diablo Range peaks, California. Res. Pap. PSW-110. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 50 p. 
50. Griffin, James R.; Critchfield, William B. 1972. The distribution of forest trees in California. Res. Pap. PSW-82. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 118 p. 
51. Gurung, Janita; Adams, A. B.; Raphael, Martin G. 1999. A review of the use of Pacific madrone by nesting, pollinating and frugivorous birds. In: Adams, A. B.; Hamilton, Clement W., eds. The decline of the Pacific madrone (Arbutus menziesii Pursh): Current theory and research directions: Proceedings of the symposium; 1995 April 28; Seattle, WA. Seattle, WA: Save Magnolia's Madrones, Center for Urban Horticulture, Ecosystems Database Development and Research: 25-32. 
52. Hagar, Donald C. 1960. The interrelationships of logging, birds, and timber regeneration in the Douglas-fir region of northwestern California. Ecology. 41(1): 116-125. 
53. Hall, Frederick C. 1974. Key to some common forest-zone plants of northwestern Washington. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 34 p. 
54. Hall, Frederick C. 1974. Prediction of plant community development and its use in management. In: Black, Hugh C., ed. Wildlife and forest management in the Pacific Northwest: Proceedings of a symposium; 1973 September 11-12; Corvallis, OR. Corvallis, OR: Oregon State University, School of Forestry, Forest Research Laboratory: 113-119. 
55. Halpern, Charles B.; Spies, Thomas A. 1995. Plant species diversity in natural and managed forests of the Pacific Northwest. Ecological Applications. 5(4): 913-934. 
56. Hamel, Dennis R. 1981. Forest management chemicals: A guide to use when considering pesticides for forest management. Agric. Handb. 585. Washington, DC: U.S. Department of Agriculture, Forest Service. 512 p. 
57. Hann, Wendel; Havlina, Doug; Shlisky, Ayn; [and others]. 2005. Interagency fire regime condition class guidebook. Version 1.2, [Online]. In: Interagency fire regime condition class website. U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior; The Nature Conservancy; Systems for Environmental Management (Producer). Variously paginated [+ appendices]. Available: http://www.frcc.gov/docs/220.127.116.11/Complete_Guidebook_V1.2.pdf [2007, May 23]. 
58. Harrington, Constance A.; Lodding, Cynthia C.; Kraft, Joseph M. 1999. Extraction and germination of Pacific madrone seed. In: Rose, Robin; Haase, Diane L., eds. Native plants: propagating and planting: Symposium proceedings; 1998 December 9-10; [Location unknown]. Corvallis, OR: Oregon State University, College of Forestry, Nursery Technology Cooperative: 38-42. 
59. Harrington, Timothy B.; Tappeiner, John C.; Walstad, John D. 1984. Predicting leaf area and biomass of 1- to 6-year old tanoak and Pacific madrone sprout clumps in southwestern Oregon. Canadian Journal of Forest Research. 14: 209-213. 
60. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. 
61. Hitchcock, C. Leo; Cronquist, Arthur. 1973. Flora of the Pacific Northwest. Seattle, WA: University of Washington Press. 730 p. 
62. Hitchcock, C. Leo; Cronquist, Arthur; Ownbey, Marion. 1959. Vascular plants of the Pacific Northwest. Part 4: Ericaceae through Campanulaceae. Seattle, WA: University of Washington Press. 510 p. 
63. Hosie, R. C. 1969. Native trees of Canada. 7th ed. Ottawa, ON: Canadian Forestry Service, Department of Fisheries and Forestry. 380 p. 
64. Hunter, John C. 1997. Fourteen years of change in two old-growth Pseudotsuga-Lithocarpus forests in northern California. Journal of the Torrey Botanical Society. 124(4): 273-279. 
65. James, Susanne. 1984. Lignotubers and burls--their structure, function and ecological significance in Mediterranean ecosystems. Botanical Review. 50(3): 225-266. 
66. Jensen, Edward C.; Anderson, Debra J. 1995. The reproductive ecology of broadleaved trees and shrubs: an overview. Corvallis, OR; Oregon State University, College of Forestry, Forest Research Laboratory. 9 p. 
67. Kartesz, John T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. 1st ed. In: Kartesz, John T.; Meacham, Christopher A. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Chapel Hill, NC: North Carolina Botanical Garden (Producer). In cooperation with: The Nature Conservancy; U.S. Department of Agriculture, Natural Resources Conservation Service; U.S. Department of the Interior, Fish and Wildlife Service. 
68. Kauffman, J. Boone; Martin, R. E. 1985. A preliminary investigation on the feasibility of preharvest prescribed burning for shrub control. In: Proceedings, 6th annual forestry vegetation management conference; 1984 November 1-2; Redding, CA. Redding, CA: Forest Vegetation Management Conference: 89-114. 
69. Kauffman, John Boone. 1986. The ecological response of the shrub component to prescribed burning in mixed conifer ecosystems. Berkeley, CA: University of California. 235 p. Dissertation. 
70. Kay, Burgess L.; Leonard, Oliver A.; Street, James E. 1961. Control of madrone and tanoak stump sprouting. Weeds. 9: 369-373. 
71. Keeley, Jon E. 1991. Seed germination and life history syndromes in the California chaparral. The Botanical Review. 57(2): 81-116. 
72. Klinka, K.; Krajina, V. J.; Ceska, A.; Scagel, A. M. 1989. Indicator plants of coastal British Columbia. Vancouver, BC: University of British Columbia Press. 288 p. 
73. Krajina, V. J. 1969. Ecology of forest trees in British Columbia. Ecology of North America. 2(1): 140-142. 
74. Krajina, V. J.; Klinka, K.; Worrall, J. 1982. Distribution and ecological characteristics of trees and shrubs of British Columbia. Vancouver, BC: University of British Columbia, Department of Botany and Faculty of Forestry. 131 p. 
75. Krcmar-Nozic, Emina; Wilson, Bill; Arthur, Louise. 2000. The potential impacts of exotic forest pests in North America: a synthesis of research. Information Report BC-X-387. Victoria, BC: Canadian Forest Service, Pacific Forestry Centre. 33 p. 
76. LANDFIRE Rapid Assessment. 2005. Reference condition modeling manual (Version 2.1), [Online]. In: LANDFIRE. Cooperative Agreement 04-CA-11132543-189. Boulder, CO: The Nature Conservancy; U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior (Producers). 72 p. Available: http://www.landfire.gov/downloadfile.php?file=RA_Modeling_Manual_v2_1.pdf [2007, May 24]. 
77. LANDFIRE Rapid Assessment. 2007. Rapid assessment reference condition models. In: LANDFIRE. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Lab; U.S. Geological Survey; The Nature Conservancy (Producers). Available: http://www.landfire.gov/models_EW.php 
78. Lotan, James E.; Alexander, Martin E.; Arno, Stephen F.; French, Richard E.; Langdon, O. Gordon; Loomis, Robert M.; Norum, Rodney A.; Rothermel, Richard C.; Schmidt, Wyman C.; van Wagtendonk, Jan. 1981. Effects of fire on flora: A state-of-knowledge review: Proceedings of the national fire effects workshop; 1978 April 10-14; Denver, CO. Gen. Tech. Rep. WO-16. Washington, DC: U.S. Department of Agriculture, Forest Service. 71 p. 
79. Maleike, Ray; Hummel, Rita L. 1999. The propagation of Arbutus menziesii from seed. In: Adams, A. B.; Hamilton, Clement W., eds. The decline of the Pacific madrone (Arbutus menziesii Pursh): Current theory and research directions: Proceedings of the symposium; 1995 April 28; Seattle, WA. Seattle, WA: Save Magnolia's Madrones, Center for Urban Horticulture, Ecosystems Database Development and Research: 94-98. 
80. Mallory, James I. 1980. Canyon live oak. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 125-126. 
81. Martin, Alexander C.; Zim, Herbert S.; Nelson, Arnold L. 1951. American wildlife and plants. New York: McGraw-Hill Book Company, Inc. 500 p. 
82. McDonald, Philip M. 1978. Silviculture-ecology of three native California hardwoods on high sites in north central California. Corvallis, OR: Oregon State University. 309 p. Dissertation. 
83. McDonald, Philip M. 1980. California black oak. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 122. 
84. McDonald, Philip M. 1980. Pacific ponderosa pine-Douglas-fir. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 120. 
85. McDonald, Philip M. 1980. Pacific ponderosa pine. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 120-121. 
86. McDonald, Philip M. 1981. Adaptations of woody shrubs. In: Hobbs, S. D.; Helgerson, O. T., eds. Reforestation of skeletal soils: Proceedings of a workshop; 1981 November 17-19; Medford, OR. Corvallis, OR: Oregon State University, Forest Research Laboratory: 21-29. 
87. McDonald, Philip M. 1992. Estimating seed crops of conifer and hardwood species. Canadian Journal of Forest Research. 22: 832-838. 
88. McDonald, Philip M.; Huber, Dean W. 1995. California's hardwood resource: managing for wildlife, water, pleasing scenery, and wood products. Gen. Tech. Rep. PSW-GTR-154. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station. 23 p. 
89. McDonald, Philip M.; Minore, Don; Atzet, Tom. 1983. Southwestern Oregon--northern California hardwoods. In: Burns, Russell M., compiler. Silvicultural systems for the major forest types of the United States. Agric. Handb. 445. Washington, DC: U.S. Department of Agriculture: 29-32. 
90. McDonald, Philip M.; Tappeiner, John C., II. 2002. California's hardwood resource: seeds, seedlings, and sprouts of three important forest-zone species. Gen. Tech. Rep. PSW-GTR-185. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station. 39 p. 
91. McDonald, Philip M. [In press]. Arbutus menziesii Pursh--Pacific madrone, [Online]. In: Bonner, Franklin T.; Nisley, Rebecca G.; Karrfait, R. P.; coords. Woody plant seed manual. Agric. Handb. 727. Washington, DC: U.S. Department of Agriculture, Forest Service (Producer). Available: http://www.nsl.fs.fed.us/wpsm/Arbutus.pdf [2007, November 9]. 
92. McDonald, Phillip M.; Tappeiner, John C., II. 1990. Arbutus menziesii Pursh Pacific madrone. In: Burns, Russell M.; Honkala, Barbara H., technical coordinators. Silvics of North America. Vol. 2--Hardwods. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service. [in press]. 
93. Menashe, Elliott. 1993. Appendix A: Plants commonly found on Puget Sound shoreland sites, [Online]. In: Vegetation Management: A program for Puget Sound bluff property owners. Publication 93-31. Olympia, WA: Washington State Department of Ecology, Shorelands and Coastal Zone Management Program (Producer). Available: http://www.ecy.wa.gov/programs/sea/pubs/93-31/app-a.html [2007, December 15]. 
94. Miller, Melanie. 2000. Fire autecology. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 9-34. 
95. Miller, Richard E.; Lavender, Denis P.; Grier, Charles C. 1976. Nutrient cycling in the Douglas-fir type--silvicultural implications. In: America's renewable resource potential--1975: The turning point; 1975 Society of American Foresters national convention; 1975 September 28 - October 2; Washington, DC: Society of American Foresters: 359-390. 
96. Minnich, Richard A.; Franco-Vizcaino, Ernesto. 1997. Mediterranean vegetation of northern Baja California. Fremontia. 25(3): 3-12. 
97. Minore, Don. 1979. Comparative autecological characteristics of northwestern tree species--a literature review. Gen. Tech. Rep. PNW-87. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 72 p. 
98. Minore, Don. 1987. Madrone duff and the natural regeneration of Douglas-fir. Res. Note PNW-RN-456. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 7 p. 
99. Morrow, P. A.; Mooney, H. A. 1974. Drought adaptations in two Californian evergreen sclerophylls. Oecologia. 15: 205-222. 
100. Munz, Philip A. 1974. A flora of southern California. Berkeley, CA: University of California Press. 1086 p. 
101. Munz, Philip A.; Keck, David D. 1973. A California flora and supplement. Berkeley, CA: University of California Press. 1905 p. 
102. O'Brien, Joseph G.; Mielke, Manfred E.; Oak, Steve; Moltzan, Bruce. 2002. Sudden oak death: Oak mortality is caused by a new pathogen, Phytophthora ramorum. Pest Alert. NA-PR-02-02. Radnor, PA: U.S. Department of Agriculture, Forest Service, State and Private Forestry, Northeastern Area. 2 p. 
103. Parker, Kathy; Hamilton, Clement W. 1999. Slope stability and Arbutus menziesii: a summary of research in Magnolia Park, Seattle, Washington. In: Adams, A. B.; Hamilton, Clement W., eds. The decline of the Pacific madrone (Arbutus menziesii Pursh): Current theory and research directions: Proceedings of the symposium; 1995 April 28; Seattle, WA. Seattle, WA: Save Magnolia's Madrones, Center for Urban Horticulture, Ecosystems Database Development and Research: 126-128. 
104. Peinado, M.; Aguirre, J. L.; Delgadillo, J. 1997. Phytosociological, bioclimatic and biogeographical classification of woody climax communities of western North America. Journal of Vegetation Science. 8: 505-528. 
105. Pelton, John. 1962. Factors influencing survival and growth of a seedling population of Arbutus menziesii in California. Madrono. 16(8): 237-256. 
106. Pojar, Jim; MacKinnon, Andy, eds. 1994. Plants of the Pacific Northwest coast: Washington, Oregon, British Columbia and Alaska. Redmond, WA: Lone Pine Publishing. 526 p. 
107. Radosevich, S. R.; Passof, P. C.; Leonard, O. A. 1976. Douglas fir release from tanoak and Pacific madrone competition. Weed Science. 24(1): 144-145. 
108. Raphael, Martin G. 1987. Use of Pacific madrone by cavity-nesting birds. In: Plumb, Timothy R.; Pillsbury, Norman H., technical coordinators. Proceedings of the symposium on multiple-use management of California's hardwood resources; 1986 November 12-14; San Luis Obispo, CA. Gen. Tech. Rep. PSW-100. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 198-202. 
109. Raphael, Martin G. 1999. Use of Arbutus menziesii by cavity-nesting birds. In: Adams, A. B.; Hamilton, Clement W., eds. The decline of the Pacific madrone (Arbutus menziesii Pursh): Current theory and research directions: Proceedings of the symposium; 1995 April 28; Seattle, WA. Seattle, WA: Save Magnolia's Madrones, Center for Urban Horticulture, Ecosystems Database Development and Research: 17-24. 
110. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. 
111. Reagan, Albert B. 1934. Plants used by the Hoh and Quileute Indians. Transactions of the Kansas Academy of Science. 37: 55-70. 
112. Reukema, Donald L. 1980. Douglas-fir-western hemlock. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 107-108. 
113. Reynolds, Hudson G.; Sampson, Arthur W. 1943. Chaparral crown sprouts as browse for deer. Journal of Wildlife Management. 7(1): 119-122. 
114. Rizzo, David M. 2003. Sudden oak death in California. In: Fosbroke, Sandra L. C.; Gottschalk, Kurt W., eds. Proceedings: U.S. Department of Agriculture interagency research forum on gypsy moth and other invasive species: 13th annual meeting; 2002 January 15-18; Annapolis, MD. Gen. Tech. Rep. NE-300. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station: 1-2. 
115. Rizzo, David M.; Garbelotto, Matteo; Davidson, Jennifer M.; Slaughter, Garey W.; Koike, Steven T. 2002. Phytophthora ramorum and sudden oak death in California: I. Host relationships. In: Standiford, Richard B.; McCreary, Douglas; Purcell, Kathryn L., tech. coords. Proceedings of the 5th symposium on oak woodlands: oaks in California's changing landscape; 2001 October 22-25; San Diego, CA. Gen. Tech. Rep. PSW-GTR-184. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 733-740. 
116. Robinson, Cyril S. 1937. Plants eaten by California mule deer on the Los Padres National Forest. Journal of Forestry. 35(3): 285-292. 
117. Rose, S. L.; Perry, D. A.; Pilz, D.; Schoenberger, M. M. 1983. Allelopathic effects of litter on the growth and colonization of mycorrhizal fungi. Journal of Chemical Ecology. 9(8): 1153-1162. 
118. Rosenberg, Kenneth V.; Raphael, Martin G. 1986. Effects of forest fragmentation on vertebrates in Douglas-fir forests. In: Verner, Jared; Morrison, Michael L.; Ralph, C. John, eds. Wildlife 2000: modeling habitat relationships of terrestrial vertebrates: Proceedings of an international symposium; 1984 October 7-11; Fallen Leaf Lake, CA. Madison, WI: The University of Wisconsin Press: 263-272. 
119. Roy, D. F. 1955. Hardwood sprout measurements in northwestern California. Forest Research Notes No. 95. Berkeley, CA: U.S. Department of Agriculture, Forest Service, California Forest and Range Experiment Station. 6 p. 
120. Roy, Douglas F. 1980. Redwood. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 109-110. 
121. Safford, Hugh Deforest. 1995. Woody vegetation and succession in the Garin Woods, Hayward Hills, Alameda County, California. Madrono. 42(4): 470-489. 
122. Sampson, Arthur W.; Jespersen, Beryl S. 1963. California range brushlands and browse plants. Berkeley, CA: University of California, Division of Agricultural Sciences, California Agricultural Experiment Station, Extension Service. 162 p. 
123. Sawyer, John O., Jr. 1980. Douglas-fir-tanoak-Pacific madrone. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 111-112. 
124. Sawyer, John O.; Thornburgh, Dale A.; Griffin, James R. 1977. Mixed evergreen forest. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley and Sons: 359-381. 
125. Smith, Walton A. 1968. The band-tailed pigeon in California. California Fish and Game. 54(1): 4-16. 
126. Stein, William A. 1980. Port Orford-cedar. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 108-109. 
127. Stein, William I. 1980. Oregon white oak. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 110-111. 
128. Stickney, Peter F. 1989. Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. FEIS workshop: Postfire regeneration. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 10 p. 
129. Stuart, John D.; Salazar, Lucy A. 2000. Fire history of white fir forests in the coastal mountains of northwestern California. Northwest Science. 74(4): 280-285. 
130. Stuart, John D.; Stephens, Scott L. 2006. North Coast bioregion. In: Sugihara, Neil G.; van Wagtendonk, Jan W.; Shaffer, Kevin E.; Fites-Kaufman, Joann; Thode, Andrea E., eds. Fire in California's ecosystems. Berkeley, CA: University of California Press: 147-169. 
131. Tappeiner, John C., II; Harrington, Timothy B.; Walstad, John D. 1984. Predicting recovery of tanoak (Lithocarpus densiflorus) and Pacific madrone (Arbutus menziesii) after cutting or burning. Weed Science. 32: 413-417. 
132. Tappeiner, John C., II; McDonald, Philip M.; Hughes, Thomas F. 1986. Survival of tanoak (Lithocarpus densiflorus) and Pacific madrone (Arbutus menziesii) seedlings in forests of southwestern Oregon. New Forests. 1: 43-55. 
133. Tappeiner, John C., II; McDonald, Philip M.; Newton, Michael; Harrington, Timothy B. 1992. Ecology of hardwoods, shrubs, and herbaceous vegetation: effects on conifer regeneration. In: Hobbs, Stephen D.; Tesch, Steven D.; Owston, Peyton W.; [and others], eds. Reforestation practices in southwestern Oregon and northern California. Corvallis, OR: Oregon State University, Forest Research Laboratory: 136-164. 
134. Tappeiner, John C. 1980. Sierra Nevada mixed conifer. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 118-119. 
135. Tarrant, Robert F. 1958. Silvical characteristics of Pacific madrone. Silvical Series No. 6. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range experiment Station. 10 p. 
136. Tehon, Leo R. 1943. Canker of Pacific dogwood and madrona. American Nurseryman. 9: 24-25. 
137. Topik, Christopher; Halverson, Nancy M.; Brockway, Dale G. 1986. Plant association and management guide for the western hemlock zone: Gifford Pinchot National Forest. R6-ECOL-230A. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 132 p. 
138. Topik, Christopher; Hemstrom, Miles A., comps. 1982. Guide to common forest-zone plants: Willamette, Mt. Hood, and Siuslaw National Forests. R6-Ecol 101-1982. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 95 p. 
139. U.S. Department of Agriculture, Natural Resources Conservation Service. 2008. PLANTS Database, [Online]. Available: http://plants.usda.gov/. 
140. Van Dersal, William R. 1938. Native woody plants of the United States, their erosion-control and wildlife values. Misc. Publ. No. 303. Washington, DC: U.S. Department of Agriculture. 362 p. 
141. Veirs, Stephen D., Jr. 1980. The role of fire in northern coast redwood forest dynamics. In: Proceedings of the conference on scientific research in the National Parks: Fire ecology; 1979 November 28 - November 28; San Francisco. Washington, DC: U.S. Department of the Interior, National Park Service: 1-20. 
142. Veirs, Stephen D., Jr. 1982. Coast redwood forest: stand dynamics, successional status, and the role of fire. In: Means, Joseph E., ed. Forest succession and stand development research in the Northwest: Proceedings of the symposium; 1981 March 26; Corvallis, OR. Corvallis, OR: Oregon State University, Forest Research Laboratory: 119-141. 
143. Volland, Leonard A.; Dell, John D. 1981. Fire effects on Pacific Northwest forest and range vegetation. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region, Range Management and Aviation and Fire Management. 23 p. 
144. Waring, R. H. 1969. Forest plants of the eastern Siskiyous: their environment and vegetational distribution. Northwest Science. 43(1): 1-17. 
145. White, Keith L. 1966. Structure and composition of foothill woodland in central coastal California. Ecology. 47(2): 229-237. 
146. Williamson, Richard L. 1980. Pacific Douglas-fir. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 106-107. 
147. Wills, Robin D.; Stuart, John D. 1994. Fire history and stand development of a Douglas-fir/hardwood forest in northern California. Northwest Science. 68(3): 205-211. 
148. Zobel, Donald B.; McKee, Arthur; Hawk, Glenn M.; Dyrness, C. T. 1976. Relationships of environment to composition, structure, and diversity of forest communities of the central western Cascades of Oregon. Ecological Monographs. 46: 135-156.