Fire Effects Information System (FEIS)

FEIS Home Page

Holodiscus discolor

• INTRODUCTORY
• DISTRIBUTION AND OCCURRENCE
• BOTANICAL AND ECOLOGICAL CHARACTERISTICS
• FIRE EFFECTS AND MANAGEMENT
• MANAGEMENT CONSIDERATIONS
• APPENDIX: FIRE REGIME TABLE
• REFERENCES

INTRODUCTORY

• AUTHORSHIP AND CITATION
• FEIS ABBREVIATION
• NRCS PLANT CODE
• COMMON NAMES
• TAXONOMY
• SYNONYMS
• LIFE FORM

The Holodiscus taxonomy is confused because oceanspray, rockspirea (H. dumosus), and small-leaved rockspirea (H. microphyllus) are taxonomically and morphologically very similar [45,163]. Authorities separating these 3 closely related taxa do so based on different leaf morphologies [45,136] and distributions [136]. This review follows the taxonomy of Lis (in [97]), who authored the Flora of North America's [63] Holodiscus chapter. In Lis's treatment, oceanspray, rockspirea, and small-leaved rockspirea are treated as separate and distinct species [97]. Some systematists lump either oceanspray and rockspirea [104,236], oceanspray and small-leaved rockspirea [103], or all 3 taxa [233] into single species.

DISTRIBUTION AND OCCURRENCE

• GENERAL DISTRIBUTION
• HABITAT TYPES AND PLANT COMMUNITIES

Conifer communities: Oceanspray is called "the most widespread and possibly the most abundant flowering shrub" in coniferous forests of northeastern Washington and northern Idaho [225]. On the Umatilla National Forest, Washington, it is dominant in coast Douglas-fir (Pseudotsuga menziesii var. menziesii), grand fir (Abies grandis), and western larch (Larix occidentalis) forests [90].

		In Oregon, coast Douglas-fir/oceanspray associations on the Willamette National Forest are primarily structurally diverse old-growth stands, containing long-lived canopy trees and a subcanopy of younger trees. Most of the stands are >150 years old [94]. Oceanspray is common to dominant in dry white fir (A. concolor) forests in the Siskiyou Mountains of southwestern Oregon [238], and it is an important shrub in Port-Orford-cedar (Chamaecyparis lawsoniana) communities of southwestern Oregon and northwestern California [257]. It is important in many mixed-conifer forests of southern Oregon and California [33,34]. These communities are codominated by Pacific ponderosa pine (P. ponderosa var. ponderosa), coast Douglas-fir, Jeffrey pine (P. jeffreyi), California black oak (Quercus kelloggii), tanoak (Lithocarpus densiflorus), and/or canyon live oak (Q. chrysolepis) [34]. Oceanspray is also important in knobcone pine (P. attenuata) communities of southern Oregon and California [39].
Oceanspray in mixed-conifer forest of Oregon's Cascade Range. Photo permission of Craig Smith.

In California, oceanspray is a characteristic to dominant species of redwood (Sequoia sempervirens) [202,254], Pacific ponderosa pine [116], shore pine (P. contorta var. contorta) [255], and Sierra Nevada lodgepole pine (P. c. var. murrayana) [77] forests. It is a minor species in pinyon-juniper (Pinus-Juniperus spp.) communities [24].

Oceanspray is associated with Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca) and Rocky Mountain lodgepole pine (Pinus contorta var. latifolia) in northern Idaho [217]; it is commonly dominant in Rocky Mountain Douglas-fir forests of northern Idaho and Montana [178,217,225]. It is infrequent to common in western redcedar-western hemlock (Tsuga heterophylla-Thuja plicata)) forests of the Interior Pacific Northwest and the Northern Rocky Mountains [148,186].

Hardwood communities: Oceanspray occurs in the understories of Oregon white oak (Q. garryana) communities throughout Oregon white oak's range ([147,185,216], review by [122]). It also occurs or dominates in other montane oak (Quercus spp.) communities in California [142].

Riparian: Oceanspray occurs in riparian communities throughout its range (review by [196]). Overstory dominants may be conifers, hardwoods or a mix [112,219]. Meriwether Lewis made the first scientific collection of oceanspray on the banks of the Clearwater River in Idaho [48]. Oceanspray is dominant in grand fir floodplain associations of eastern Washington [112]. In western hemlock stands in the central Cascade Range of Washington, it was more common on high floodplains than on low floodplains [229]. On Myrtle Island Research Natural Area, Oregon, oceanspray is occasional in red alder-Oregon ash (Alnus rubra-Fraxinus latifolia) and willow/field horsetail (Salix spp./Equisetum arvense) riparian communities [219].

Shrublands: Oceanspray is common to dominant in mixed montane shrublands of the Pacific Northwest and the Northern Rocky Mountains [51,204]. These communities are common on harsh slopes and in coniferous forests in early succession [51]. In montane regions of Nevada and western Utah, oceanspray occurs in mosaics of mountain meadow and mountain big sagebrush (Artemisia tridentata subsp. vaseyana) stands [65]. In Redwood National Park, California, it occurs in Lewis' mockorange/brittle bladderfern (Philadelphus lewisii/Cystopteris fragilis) and Sierra gooseberry/varileaf phacelia (Ribes roezlii/Phacelia heterophylla) bald-hill communities [211]. On the Jasper Ridge Biological Reserve in coastal northern California, oceanspray is an associated species in chamise (Adenostoma fasciculatum) chaparral communities [1].

Publications describing plant communities where oceanspray is a dominant or indicator species are listed below.

• western hemlock/salal (Gaultheria shallon)-oceanspray forest association of the Olympic National Forest [95]
• western hemlock-coast Douglas-fir/oceanspray association of the Gifford Pinchot National Forest [222]
• coast Douglas-fir-Pacific madrone (Arbutus menziesii)/hairy honeysuckle (Lonicera hirsuta)-oceanspray and coast Douglas-fir-Pacific madrone/salal-oceanspray associations across the Puget Trough of west-central Washington [32]
• Rocky Mountain lodgepole pine/Rocky Mountain maple-Saskatoon serviceberry (Acer glabrum-Amelanchier alnifolia)-oceanspray woodlands and forests; occur on the east side of the Cascade Range, the Okanogan Highlands, and the Blue Mountains [43]
• Pacific ponderosa pine-coast Douglas-fir/Saskatoon serviceberry-oceanspray communities of the Blue Mountains [84]
• oceanspray and oceanspray-mallow ninebark shrublands on north-facing slopes of the Blue Mountains [51]
• east-canyon mixed shrublands of the Columbia Basin and Blue Mountains [44]
• grand fir-oceanspray floodplain association of eastern Washington; this type is a variant of the grand-fir-common snowberry (Symphoricarpos albus) floodplain association [112]
• Rocky Mountain Douglas-fir-Pacific ponderosa pine-Rocky Mountain lodgepole pine/mixed shrub forest zone in the Wallowa Mountains [38]
• Pacific ponderosa pine/mallow ninebark (Physocarpus malvaceus)-oceanspray association of southeastern Washington [152]
• Rocky Mountain Douglas-fir-mallow ninebark-oceanspray habitat type of eastern Washington

• coast Douglas-fir/oceanspray forest communities in the west-central Cascade Range; characterized by open stands of old growth [57]
• coast Douglas-fir/oceanspray/grass and coast Douglas-fir/oceanspray/vine maple (Acer circinatum) community types of the western Cascade Range; the former is the most widely distributed of Oregon's dry-forest types [153]
• Rocky Mountain lodgepole pine/Rocky Mountain maple-Saskatoon serviceberry-oceanspray woodlands and forests; occur on the east side of the Cascade Range and the Blue Mountains [43]
• oceanspray and oceanspray-mallow ninebark shrublands on north-facing slopes of the Blue Mountains [51]
• Pacific ponderosa pine-Rocky Mountain Douglas-fir/western snowberry-oceanspray communities of the Blue Mountains [84]
• east-canyon mixed shrublands of the Columbia Basin and Blue Mountains [44]
• westside Oregon white oak/oceanspray and dry coast Douglas-fir/oceanspray woodlands and forests; mostly in Willamette Valley and the Klamath Mountains [33]
• coast Douglas-fir/oceanspray-whipplevine (Whipplea modesta) series on the Coast Ranges; occurs on the Medford District of BLM and on the Siskiyou and Rogue River National Forests. Indicative of dry sites [19].
• coast Douglas-fir/oceanspray/salal habitat type off the southern coast [20]

• coast live oak (Q. agrifolia)/oceanspray-common snowberry subseries of California's hardwood rangeland cover types; on upper-elevation (>1,250 feet (380 m)), mesic sites [9]
• redwood-coast Douglas-fir-hardwood/oceanspray vegetation types of the North Coast Ranges [254]
• canyon live oak (Q. chrysolepis)-oceanspray forest cover type of central California [142]

• Rocky Mountain Douglas-fir/mallow ninebark-oceanspray habitat type [51,152]
• Pacific ponderosa pine/mallow ninebark-oceanspray association

• Rocky Mountain Douglas-fir/oceanspray and Rocky Mountain Douglas-fir/mallow ninebark-oceanspray forest habitat types [178]
• climax western redcedar-western hemlock, Rocky Mountain Douglas-fir, grand fir, and subalpine fir (A. lasiocarpa)-Rocky Mountain Douglas-fir-grand fir forests [186]
• indicator species of the Rocky Mountain Douglas-fir/mallow ninebark habitat type [125]

• indicator species of dry western redcedar-western hemlock/shrub, Pacific ponderosa pine/shrub, and Rocky Mountain Douglas-fir/shrub forests [148]

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

• GENERAL BOTANICAL CHARACTERISTICS
• SEASONAL DEVELOPMENT
• REGENERATION PROCESSES
• SITE CHARACTERISTICS
• SUCCESSIONAL STATUS

Botanical description Raunkiaer life form Botanical description: This description covers characteristics that may be relevant to fire ecology and is not meant for identification. Keys for identification are available (for example, [97,99,117,193]). Morris and others [161] provide a key for identifying oceanspray and other shrubs in winter. Oceanspray and rockspirea are distinguished by their forms, leaf characteristics, and distributions [136]; intergradation of the 2 species is most pronounced in Nevada [104] and Utah [236].
		Photo © 2009 Barry Breckling

Morphology:
Form: Oceanspray is a deciduous [1,41,198], spreading shrub with slender arching branches [75,136,223]. It can range from bushy forms about 2.5 feet (0.75 m) tall on poor or frequently disturbed sites to arborescent forms that may be 20 feet (6.1 m) tall in coastal areas. Plants are usually 3 to 10 feet (1-3 m) in height [52,99,198]. They typically have multiple branches [198]. Stem wood is hard and dense [73]; bark of mature plants is shreddy [41,174]. Stand structures of plant communities where oceanspray is important are discussed in the Stand structure section of Fuels.

Leaves and flowers: The leaves are mostly 1.6 to 2.75 inches (4-7 cm) long and 0.8 to 2.75 inches (2-7 cm) wide [136,223]. Oceanspray has a large leaf area relative to most associated shrubs. In the Siskiyou Mountains of southwestern Oregon, its leaves were more densely packed, larger, thinner, and more prone to wilt than leaves of associated shrub species [40].

Oceanspray flowers are small, about 2 mm long [163]. They are borne on large, showy, terminal panicles that may reach 12 inches (30 cm) long [22]. The name "oceanspray" is derived from these masses of loose, creamy plumes [50]. The fruit is a 1-seeded [45] achene [41,97,117], about 2 mm long [198].

Roots: Rooting depth is likely associated with depth to bedrock. In southwestern Oregon, oceanspray extracted water from no deeper than 3 feet (1 m) below ground, indicating a shallow root system [40]. Dyrness and Franklin [57] had similar findings in the west-central portion of the Cascade Range in Oregon, where shallow soils confined roots to <3 feet below the soil surface. However, a planting guide for the Pacific Northwest reports oceanspray roots as "deep and wide" [154], and researchers described oceanspray as "relatively deep-rooted" in the Blue Mountains [249].

Descriptions of oceanspray's root morphology were not found in the literature as of 2010.

Life span: This species rarely lives more than 30 years [10]. On the Jasper Ridge Biological Reserve, California, its mean life span was 4.5 years [1].

Physiology: Oceanspray is highly drought tolerant [51]. It has adapted to dry sites and drought by shutting down or slowing its rate of transpiration. In droughty conditions, it apparently uses water less efficiently than associated sclerophyllous species. Its large leaf area, however, may partially compensate for low water transpiration rates in summer. It is likely that oceanspray depletes water in upper soil layers rapidly in summer [40].

Antieau [13] suggested that oceanspray may differ in water-use efficiency and cold tolerance across its distribution.

Leaf phenology is closely regulated by weather. In the Siskiyou Mountains of Oregon, leaf water conductance peaked in July [40]. On the Jasper Ridge Biological Preserve, mean leaf age was 4.5 months; leaves were drought-deciduous and mostly absent by August [1]. A study in the western redcedar-western hemlock zone of northern Idaho found summer or fall drought initiated leaf color change and leaf drop [55].

Oceanspray consistently shows a late and long period of flowering throughout its distribution:

Pollination and breeding system: Insects pollinate oceanspray (review by [196]). The flowers are perfect [100].

Antieau [13] suggested that mountain ranges restrict oceanspray breeding. A study across oceanspray's distribution in Washington and Oregon showed phenotypic differences in oceanspray (for example, in leaf area); these differences were related to geographic regions and climate [13].

Flower and seed production: Thinning, burning, or other canopy-opening events may increase oceanspray's seed output. In the understories of coast Douglas-fir forests in western Oregon, oceanspray showed a "large increase" in flower and/or fruit production after moderate thinnings (leaving 200 trees/ha) or heavy thinnings (leaving 100 trees/ha or 0.4-ha openings); production increases were "minimal" after light thinning (leaving 300 trees/ha) [213].

Seed dispersal: Oceanspray seed is disseminated by wind ([213,237,248], review by [198]) or animals [213].

Seed banking: Oceanspray has a soil seed bank [60,150]. In western hemlock forests of southwestern British Columbia, viable oceanspray seed was more common in undisturbed soils (x=15.5 germinants/0.04-m² soil sample) compared to clearcut rights-of-way in early-seral succession (x=1-2.5 germinants/0.04-m² soil sample) [150].

Germination, seedling establishment, and plant growth: Fresh oceanspray seed is dormant [74]. In the field, it likely requires overwintering to germinate. As of 2010, little research had been conducted on oceanspray's germination requirements [197]. Stratification at around 41 ^oF (5 ^oC) [197] for 15 to 18 weeks breaks dormancy in the laboratory [12,113,197].

Oceanspray seed may have low viability. According to a fact sheet, most seeds lack developed embryos, so only about 7% of a given seed lot may be sound [74].

Seedling establishment is uncommon [140,149,150,208,213] but has been documented a few times. Open stand structure [214], heat, and bare mineral soil may favor oceanspray germination and establishment (review by [196]). In the Oregon Coast Range, oceanspray seedlings emerged well (>70%) in both clearcuts and young, unthinned conifer stands; however, seedlings survived only in the young, unthinned stands [214]. Oceanspray established from seed 3 growing seasons after a debris flow on the Central Coast Ranges of southwestern Oregon [173]. See Seedling establishment in the Plant Response to Fire section for studies on postfire seedling establishment.

A review states that oceanspray seedlings grow slowly in their first 2 years of development [196]. Plants released by overstory removal may grow rapidly, however. Daubenmire and Daubenmire [51] found that in northern Idaho, oceanspray grew up to 15 feet (4.6 m) tall following harvest of the Rocky Mountain Douglas-fir overstory; this was twice its stature in unharvested Rocky Mountain Douglas-fir forests.

Oceanspray is common in sands and clay loams (review by [196]) but may occur in all soil textures. In the Cascade Range of Oregon, coast Douglas-fir/oceanspray communities occur on coarse soils and loams but not on fine soils [153]. A study in the Blue Mountains, however, found oceanspray presence was positively correlated with fine-textured soils (P<0.05) [249]. In western redcedar-western hemlock forests of northern Idaho, oceanspray cover, frequency, and importance value increased as soil organic matter increased; increases in importance values were significant (P=0.05) [165]. Soils supporting oceanspray are often stony [67,87,94,97,98,100], and oceanspray sometimes grows within rock crevices. It is common on talus slopes (review by [196]). In Nevada and western Utah, oceanspray grew on talus slopes near mountain meadows and in granite boulder piles [65].

Oceanspray occurs on a variety of parent materials. In the Cascade Range of Oregon, coast Douglas-fir/oceanspray communities occur on poorly developed basalts, andesites, and other parent materials of volcanic origin [153]. At Oregon Caves National Monument, mixed-conifer forests with oceanspray occur on soils of diorite origin [240]. Poison-oak (Toxicodendron diversilobum)-oceanspray-Mexican elderberry (Sambucus mexicana) communities of San Luis Obispo County, California, are associated with andesite-derived soils [235].

Photo © Br. Alfred Brousseau, St Mary's College

Moisture regime: Oceanspray is most common on dry sites. McDonald and others [148] list oceanspray as an indicator species of dry montane/shrub forests of the Northern Rocky Mountains. Oceanspray is also associated with dry montane forests in British Columbia [168,179] and elsewhere in the Pacific Northwest. It is an indicator species of very dry to moderately dry, nitrogen-medium soils in coastal British Columbia [110,111]; its occurrence decreases with increasing precipitation [111]. It also grows in dry to fresh soils in coniferous forests of interior British Columbia [168]. The western hemlock-coast Douglas-fir/oceanspray association occurs on some of the hottest and driest sites in the Cascade Range of Washington [222]. In the west-central portion of the Cascade Range in Oregon, Dyrness and Franklin [57] found the coast Douglas-fir/oceanspray association occurs on the dry end of coast Douglas-fir forest types. In an extreme case, oceanspray is "widespread but not abundant" on the Indian Plateau of southwestern Oregon. The plateau is a severe site known for widely fluctuating and extreme temperatures in winter and summer and a record of poor artificial regeneration of conifers [156].

Oceanspray is also reported from sites with moist to mesic soils. It is frequently associated with riparian communities (review by [196]). In southeastern Washington and northern Idaho, Pacific ponderosa pine/mallow ninebark-oceanspray communities dry out later in the growing season than Pacific ponderosa pine/common snowberry communities [152]. Oceanspray occurs on moist woodland edges in California [97] and in moist open woods in British Columbia [193]. In western redcedar-western hemlock forests of northern Idaho, oceanspray frequency was significantly greater on sites with 21% to 25% soil moisture content than on sites with drier or wetter soils [165].

Aspect and topography: This species is most common on warm, dry, south-facing slopes [67,98,165]. A grand fir/oceanspray association in southwestern Washington is common on exposed, south-facing slopes and on ridgetops. Sites having this association remain snow-free much of the year and experience extreme summer drought [221]. In Douglas-fir (Pseudotsuga menziesii) forests in the Columbia River Gorge of Washington, oceanspray had greatest cover on south-facing slopes (13%) and least cover in mesic ravines (6%) [243]. The coast Douglas-fir/oceanspray association in Oregon's Coast Ranges occurs most often on relatively steep, south- or west-facing slopes between 2,000 and 3,000 feet (600-900 m) elevation. The environment is hot and dry, and the growing season is long, with drought developing by midsummer. Snowpacks are not generally deep or persistent [94]. The western hemlock-Douglas-fir/oceanspray association is found in some of the hottest and driest forests in the western Cascade Range. Sites are "always" upper slopes and fairly steep, and drainage and solar input are "excessive" [87]. Coast Douglas-fir/oceanspray communities in the Cascade Range of Oregon are also most prevalent on dry, south-facing slopes [153]. In western redcedar-western hemlock forests of northern Idaho, oceanspray cover, frequency, and importance values were significantly greater on south- than north-facing slopes (P=0.05) [165].

Oceanspray grows on more mesic exposures as well. In montane zones on the Umatilla National Forest, it dominated the understory on north, northeast, northwest, and east aspects [90]; in western Oregon clearcuts it occurred only on north-facing slopes [246]. Oceanspray mostly grows on moist slopes in southern California [41], where it reaches the southern end of its distribution.

Elevation: Oceanspray occurs from sea level to about 7,000 feet (2,150 m) across its range. It mostly grows on low-elevation montane sites. In western redcedar-western hemlock forests of northern Idaho, oceanspray cover, frequency, and importance values were significantly greater on 3,000- to 3,400-foot (910-1,000 m) elevations than on higher-elevation sites (P=0.05) [165]. Oceanspray grows mostly on high peaks in the Great Basin (review by [196]).

Climate: Oceanspray occurs mostly in dry zones [135], although it is characterized as a "predominantly humid zone species" in western Washington [53]. Annual precipitation across its United States distribution [37,56,97,137,215,241] ranges from 9.3 inches (236 mm) in central Oregon [56] to 57 inches (1,140 mm) in western Washington [137].

Seral occurrence: Disturbance favors oceanspray [111,113]. Kruckeberg [113] characterized oceanspray as a "colorful reclaimer of open or disturbed lands" of the Pacific Northwest, where it commonly establishes on recently logged sites, in second growth, and on roadbanks. It is especially common in seral Douglas-fir forests [111]. Following the Sundance Fire in northern Idaho, oceanspray was important or codominant in the first decade of postfire succession in Rocky Mountain Douglas-fir-western hemlock forests [206]. On another site in northern Idaho, oceanspray grew rapidly and dominated early-seral sites after a Rocky Mountain Douglas-fir forest was clearcut. The shrub layer regained precutting cover about 60 to 80 years after tree harvest [51]. Oceanspray seedlings established 3 growing seasons after a debris flow on the Central Coast Ranges of southwestern Oregon [173].

Oceanspray prefers open sites [111,218]. It is described as a "light demanding, early successional" species [218]. Logging and fire promote oceanspray by opening the canopy. A study at the Eastern Oregon Experiment Station showed shrub cover, including that of oceanspray, decreased with increasing cover of the mixed-conifer overstory. At about 90% canopy closure, shrub cover dropped to about 5%. However, even under a nearly closed canopy, a few shrubs remained alive in the understory, and seedlings of these shrubs established in canopy breaks [247]. In coast Douglas-fir/salal stands on foothills of the Cascade Range, Washington, maximum oceanspray cover occurred approximately 20 years after disturbance (clearcutting or wildfire); oceanspray generally declined after that [137]:

Defoliation and/or death of overstory trees due to insects may favor oceanspray. In the Blue Mountains, oceanspray showed 5% cover and 15% frequency 23 years after a record-breaking, 2-year attack by Douglas-fir tussock moths. About 1,250 miles² (3,240 km²) of a grand fir-Douglas-fir forest was affected by the outbreak [250].

Where it is a minor species, oceanspray may not decline with canopy closure. In western redcedar-western hemlock forests of northern Idaho, its cover, frequency, and importance values were not significantly different in 5 canopy-cover classes ranging from 1% to 100% closure. Oceanspray had ≤1% cover in all canopy-cover classes. Similarly, its cover, frequency, and importance values in these forests were not significantly different between logged, logged-and-burned, single-broadcast-burned, or multiple-broadcast-burned sites and sites with no history of logging or prescribed fire [165].

Logging: Lightly-shaded areas, such as those occurring a few decades after thinning, can promote oceanspray growth [194]. In Douglas fir-western hemlock forests of coastal Oregon, oceanspray was associated with intermediate tree densities (P<=0.01) [199]. In Douglas- fir stands in northern Idaho, its cover peaked about 20 years after logging [176].

On the Fort Lewis Military Reservation of Washington, a late 1990s study found oceanspray cover was greater in a coast Douglas-fir/oceanspray forest that had been clearcut in the 1920s and thinned twice afterwards (2.5% oceanspray cover) than in a coast Douglas-fir/oceanspray forest that had been partially cut only once, in the 1930s (1.5% oceanspray cover) [220]. In Pacific ponderosa pine and Rocky Mountain Douglas-fir habitat types of the Swan Valley, Montana, oceanspray cover was greater on clearcut (15%) and plantation (10%) plots than on untreated plots (8%) [69].

In the Klamath Mountains of Oregon and California, shrubfields of oceanspray and other sprouting shrubs develop after logging or fire when conifers fail to regenerate in the early postfire community; conifers eventually replace the shrubs on most sites [149].

Logging does not favor oceanspray on all sites. In northern Idaho logging reduced oceanspray frequency slightly compared to its frequency in the understory of an adjacent unlogged site. The study site was in a western hemlock/pachistima forest. A tall-shrub (>3 feet (1 m)) community developed after logging; oceanspray was a component of this early-seral, tall-shrub community. On cut sites, oceanspray had 1.4% frequency 7 years after logging and 0.7% frequency 25 years after logging. It had 2.1% cover on the unlogged site 25 years after treatments [242].

See Plant response to fire for more information on oceanspray occurrence in seral postfire communities.

FIRE EFFECTS AND MANAGEMENT

• FIRE EFFECTS
• FUELS AND FIRE REGIMES
• FIRE MANAGEMENT CONSIDERATIONS

FIRE EFFECTS:
Immediate fire effect on plant: Oceanspray is usually top-killed by fire [25,35,60,140,167,167,203,207,207,253]. Many fire researchers describe it as "moderately resistant" to fire [60,61,207,248]. Stickney [208] classifies oceanspray among the shrubs "least susceptible" to fire mortality, although fire may kill some individuals within a population [91].

Postfire regeneration strategy [207]:
Tall shrub, sprouting root crown
Small shrub, sprouting root crown
Ground residual colonizer (on site, initial community)
Initial off-site colonizer (off site, initial community)
Secondary colonizer (on- or off-site seed sources)

Fire adaptations and plant response to fire:
Fire adaptations: Oceanspray sprouts from the root crown after fire [55,60,61,140,167,208,209,228,232,245]. Stickney [207,208] uses oceanspray as a characteristic example of a root crown-sprouting, "postfire survivor species".

Oceanspray may establish from soil-stored seed [25,60,160,167,207,232,248], although seedlign establishment is apparently rare [167,248] and to date (2010), has only been documented 3 times after fire [83,160,191]. Postfire seed dispersal onto burns by wind or animals is possible but has not been documented. Heat and bare mineral soil may favor oceanspray germination and establishment (review by [196]); this has not been determined experimentally or in the field.

Plant response to fire: Oceanspray sprouts from the root crown and establishes from seed after fire.

Sprouting from surviving root crowns is oceanspray's most common method of postfire regeneration [25,25,35,42,54,58,108,126,134,167,167,206,207,208,208,232,245,248,248]. It usually takes 5 to 10 years for oceanspray to recover its prefire abundance [55]. It is often important on early-seral shrubfields after fire; these shrubfields eventually succeed to coniferous forests [204]. Shatford and others [195] found oceanspray "common and abundant" on 9- to 19-year-old burns in dry coast Douglas-fir forests of the Siskiyou and Klamath mountains of Oregon and California. Oceanspray's cover generally increases after disturbances [25,35,167,248] that open the canopy in late-successional forests, so oceanspray is likely to increase on sites that were in late succession before fire [42,60,61,167,248].

Sprouting allows oceanspray to survive large, severe wildfires. In southeastern Oregon, oceanspray was noted in montane postfire communities after the 2002 Biscuit Fire Complex, a series of mixed-severity (low- to high-severity surface and crown) wildfires that burned about 500,000 acres (200,000 ha), and after the Bear Butte and Booth Fire Complex, a series of mixed-severity wildfires that burned about 91,000 acres (37,000 ha) [85]. A survey of wildfire-burned sites in California chaparral and moist and dry, mixed-conifer forests of western Montana showed oceanspray sprouted after fires of all severities in all vegetation types where it was present before fire. Its postfire sprouting response was stronger in moist and dry conifer forests than in chaparral [134]. Reburns generally favor oceanspray (see Northern Idaho below).

Seedling establishment: Oceanspray regenerates from soil-stored seed after fire [25] occasionally. Neuenschwander [167] noted that for oceanspray, "reproduction from seed is rarely observed after a burn", and Stickney (1978 personal communication cited in [167]) noted that oceanspray seedling growth is slow in burns compared to other species. However, in the western Cascade Range of Oregon, oceanspray seedlings occurred 2 years after former western-hemlock/Douglas-fir old growth had been clearcut, broadcast burned, and planted to coast Douglas-fir. Oceanspray was not present on study plots before treatments [191].

Although oceanspray does not often establish from seed, its postfire seedling cover may be greatest where fire was severe. A northern Idaho study determined that oceanspray regenerated from duff- and soil-stored seed 1 to 3 years after broadcast burning in western redcedar/queencup beadlily (Clintonia uniflora) habitat types. Greatest oceanspray seedling cover was in areas that experienced severe fire [160]. Succession proceeded slowly after the severe 1960 Saddle Mountain Fire in the Saddle Mountain Wilderness, Arizona, where oceanspray is near its southern distributional limit. Oceanspray showed low cover (0.03%) and frequency (0.01 %) in a vegetation survey conducted in postfire year 48 [83].

Pacific Northwest:
Oceanspray showed good recovery after prescribed-burning and thinning-and-burning treatments in the Pacific Northwest. Herbicides-and-burning treatments helped control oceanspray.

Washington: In a short-term study, oceanspray gained cover rapidly between postfire years 2 and 3 following a July 1970 wildfire in North Cascades National Park. The wildfire burned a coast Douglas-fir-western hemlock stand on a ridgetop [155].

Near Wenatchee, oceanspray was abundant a year after thinning and prescribed fire in a Pacific ponderosa pine/white spirea (Spiraea betulifolia) forest. Burning was conducted on 22 September 1997; the fire was of low severity, with flame lengths generally <4.9 feet (1.5 m). Seventy-eight percent of oceanspray plants sprouted in postfire year 1. Ocean spray density was 15 plants/ha in postfire year 1; 337 plants/ha in postfire year 3; and 238 plants/ha in postfire year 9. The management objectives—to reduce fuels and conifer seedling density and to help restore approximate historical stand structure and composition—were considered met [91].

Oceanspray had regained its "former size and luxuriance" 10 years after slash burning in a Pacific ponderosa pine/mallow ninebark habitat type in northeastern Washington [232].

Central Oregon:
Spraying and prescribed burning for shrub control: In the central Oregon coast, an herbicides-and-fire treatment helped control oceanspray in the short term. A shrubfield on a heavily logged Sitka spruce (Picea sitchensis)-western hemlock habitat type was treated with either herbicides (picloram-phenoxy mix), herbicides followed by October prescribed fire, herbicides followed by tractor crushing, or bulldozer scarification without herbicides or fire. The objective was to reduced shrub interference with conifer seedlings. For sprayed plots, herbicide was applied in summer or early fall; any additional treatments were done in fall. Treatments employing mechanical disturbance were most effective, and spraying alone least effective, at controlling oceanspray. The author attributed the only moderate control obtained from the herbicides-and-fire treatment to recent rains and poor combustion of fuels [107].

Although herbicides and crushing best controlled oceanspray in particular, scarification controlled shrubs in general better than other treatments [107].

Southern California: Prescribed burning had little or no effect on oceanspray in a mixed-conifer Jeffrey pine-California black oak woodland with a chaparral understory in Cuyamaca Rancho State Park, San Diego County. In postfire year 2, oceanspray was present in trace amounts (4 plants/ha) on unburned control plots but was not present on burned plots [143]. See the Research Project Summary of Martin and Lathrop's [124,143,144] study for details of the fire weather, fire prescription, and fire's effects on 38 other plants in the woodland.

Northern Rocky Mountains:
In the Northern Rocky Mountains, prescribed fires had either no effect or a positive effect on oceanspray. Oceanspray had a strong postfire recovery after a wildfire. Prescribed and wildfire reburns promoted oceanspray and other shrubs over conifers. Postfire ungulate browsing of oceanspray was minimal.

Western Montana: Thinning and/or prescribed fire had little effect on oceanspray in Pacific ponderosa pine and Rocky Mountain Douglas-fir forests on the Lubrecht Experimental Forest. Oceanspray was scarce in these forests; it had <1% cover and frequency before treatments and in posttreatment year 2. The treatments successfully reduced fuel loads and helped restore historical stand structure in these forests. See Metlen and others' Research Project Summary for information on fire and thinning prescriptions used in this study, on fire behavior, and on the effects of thinning-alone and/or thinning-and-burning treatments on 7 tree, 24 shrub, 37 graminoid, 1 fern, and 540 forb species.

Northern Idaho: In Rocky Mountain Douglas-fir/mallow ninebark habitat types on the Coeur d'Alene Reservation, mean oceanspray across 3 postfire years cover did not differ significantly on unburned control sites, sites burned at low intensity (127 Kcal/m-s), or sites burned at high intensity (781 Kcal/m-s). For information on the fire weather, fire prescription, and postfire responses of 14 other shrub, 9 perennial graminoid, 25 perennial forb, and 6 annual species, see the Research Project Summary of Armour and others' [15] study.

Oceanspray showed mixed but mostly favorable responses after prescribed fires on the Clearwater National Forest [18]. Prefire abundance and season of burning were not given.

Oceanspray grew rapidly after prescribed fires in seral shrubfields in a grand fir/pachistima (Pachistima myrsinites) habitat type on the Coeur d’Alene National Forest. Its sprouts were approximately 24 inches (60 cm) tall by the end of the 1st postfire growing season. Spring-burned oceanspray sprouted during postfire weeks 4 to 8; plants burned in fall did not sprout until the next spring. Unlike many associated shrubs that sprouted most prolifically following spring prescribed fire, oceanspray showed greatest shoot production following fall prescribed fire [133].

Ten years after the mixed-severity Sundance Fire, Stickney [206] reported 5% to 11% cover of oceanspray; it was one of the principal cover species on his plots. Study plots were in a western hemlock habitat type. Oceanspray was a component of the initial "sprouting shrub stage" that formed a year after the fire and persisted through at least 10 years of postfire succession [206].

Reburning: Reburns generally promote oceanspray and other sprouting shrubs over conifers. In conifer forests on the Coeur d’Alene National Forest, oceanspray was a component of mixed-species shrubfields after wildfires 19, 21, and 9 years apart (initial fire in 1851, reburns in 1870, 1910, and 1919). The shrubs were dense and well developed 10 years after the 1919 wildfire [123].

Oceanspray sprouted and grew rapidly after a March 1965 prescribed fire in a grand fir/Oregon boxwood (Paxistima myrsinites) habitat type in the Fish Creek area of the Nez Perce National Forest. Prior to the prescribed fire, the overstory had been killed by wildfires in 1934 and in 1954; a shrubfield developed after these fires. The fire management objective was to maintain the shrubfield as big game browse [131,132].

The fire successfully rejuvenated the shrubfield. See the 2nd Fire Case Study in Saskatoon serviceberry for details on the prescription and behavior of this prescribed fire.

Prescribed fire and postfire browsing: Since oceanspray is relatively unpalatable, it is unlikely to be overbrowsed after fire. Ungulates browse new oceanspray sprouts, but browsing pressure usually declines rapidly after postfire year 1. On the Sherman Creek Watershed in northern Idaho, elk browsed new oceanspray sprouts but did not consume unburned plants or sprouts ≥3 years old. Prescribed burning was conducted in the spring or fall of 1966 to enhance winter browse for big game species, so fall-burned plants did not provide forage until the next growing season (1968). Oceanspray was not utilized in postfire years 6 or 8 on either burned or unburned plots; overall, elk use of oceanspray was the least of 8 browse species. Diameter of oceanspray twigs that sprouted after fire was thicker than that of twigs of unburned plants for at least 2 postfire years [128]:

Prescribed fire and cattle-grazing treatments had no significant effect on oceanspray cover in a Rocky Mountain Douglas-fir/mallow ninebark habitat type on the Idaho Experimental Forest [253].

See Palatability for more information on ungulate use of oceanspray after fire.

Fuels Fire regimes
	Photo by J. E. (Jed) and Bonnie McClellan © California Academy of Sciences

Fuels: Oceanspray is most frequent on south slopes of dry montane forests (see Site Characteristics); these sites typically burn earlier in the season or with higher severities than cooler, drier sites. In northern Idaho, Smith and Fischer [204] placed the Pacific ponderosa pine, Rocky Mountain Douglas-fir, and grand fir forests where oceanspray is most typically dominant in Fire Group 2. These forests tend to have warm temperature regimes, dry to moderate soil moisture, and are generally more productive—with heavier loads of downed woody fuels—compared to cooler or drier forests. Oceanspray also occurs in mesic to moist grand fir forests (Fire Group 7); these forests also have heavy fuel loads. See Smith and Fischer [204] for fuel load measurements representative of coniferous forest habitat types where oceanspray is important in northern Idaho.

In Pacific ponderosa pine-grand fir forests of Washington and Oregon, growing-season moisture content of shrubs, including oceanspray, averaged >125% over 2 years. Shrub moisture content peaked in June at ~175%. Moisture contents of dominant overstory trees are also described in this study [7].

In many forest types with oceanspray, fire exclusion has resulted in higher loads of woody debris compared to woody fuel loads when historic fire regimes were still functioning. In white fir stands in the Siskiyou Mountains of southwestern Oregon, large woody-debris loads were positively correlated with time-since-fire (P=0.01). Snag density was positively correlated with low (30 snags/ha) and high (23 snags/ha) fire severities (P=0.05). The authors attributed the correlation to nonconsumption of preexisting snags at low fire severity and creation of new snags at high severity. In this study, oceanspray dominated the shrub layer of white fir stands in dry, interior valleys [238].

Stand structure: Stand structure of communities where oceanspray is an important component of the vegetation is variable, as is the amount of fuel oceanspray contributes. On some sites, structure is open, with a sparse shrub component. In the west-central portion of the Cascade Range of Oregon, the coast Douglas-fir/oceanspray association displayed a relatively open stand structure (30-60% crown closure) of old growth, with few shrubs and a "very poorly developed" herb layer. Oceanspray cover averaged 5% [57]. Another study of coast Douglas-fir/oceanspray communities in the Cascade Range found that except for oceanspray, the tall-shrub layer was depauperate; low-shrub and herbaceous cover was also low. Incense-cedar, however, was encroaching in the subcanopy. Tree densities averaged 53 stems/ha for coast Douglas-fir and 5/ha for incense-cedar [153]. Live shrubs, including oceanspray as a dominant, comprised <5% of total stand biomass in mixed-conifer communities in the White Cap Wilderness Study Area of northern Idaho. Total shrub fuel loads ranged from 204 to 2,190 lbs/acre; shrubs were 0.6 inch to 27 inches (1.5-69 cm) tall, with 2% to 50% cover [26].

Some communities with oceanspray have denser overstories and/or understories. A coast Douglas-fir/oceanspray stand on the west-central portion of the Cascade Range, Oregon, had 70% tree cover, 46% shrub cover, and 36% herb cover. Aspect of the forest was southwest; it was the hottest and driest of 18 stand types examined [256]. Bailey [20] found coast Douglas-fir/oceanspray-salal habitat types off the southern coast of Oregon had relatively open canopies and "well-developed" shrub layers. Oceanspray averaged 30% cover [20]. Western redcedar-western hemlock forests often have a dense overstory, but understory cover of oceanspray and other shrubs may be sparse [204].

Mixed-conifer forests of southern Oregon and California are structurally and compositionally complex, with small conifers—often white fir and/or incense-cedar—often forming ladder fuels in a well-developed subcanopy. Snags and large, downed woody debris are common, but fuel loads are highly variable. Many of these mixed-conifer forests support a moderate to dense shrub understory, although some have few shrubs but a dominant herbaceous layer, and others have both depauperate understory and ground layers [34]. Stand structure in California's mixed-conifer forests was mostly open in the presettlement period [79].

Insect attacks increase snag densities in oceanspray habitats, which eventually increase dead and downed woody fuel loads. Youngblood and Wickman [250] provide data on stand structure, live and dead tree abundance, and shrub and herb cover of a grand fir-Douglas-fir forest attacked by Douglas-fir tussock moths 23 years prior. Oceanspray was an important component of the understory (5% cover); the site was in the Wenaha-Tucannon Wilderness in the Blue Mountains of Washington and Oregon [250].

Models: A few models were available for predicting oceanspray's contribution to total fuel loads as of 2010. Smith and Brand [205] review equations for predicting oceanspray biomass. Harris [90] presents models to predict oceanspray aboveground biomass and cover; the models were developed from data collected in coast Douglas-fir, grand fir, and western larch forests on the Umatilla National Forest. Brown [27] provides a model for predicting total aboveground oceanspray biomass and total leaf biomass based on basal stem diameter. Samples on which the model is based were collected in northern Idaho and western Montana [27].

Leaf area indices are used in some fuel models [49]. In the Siskiyou Mountains of southwestern Oregon, oceanspray had a large mean leaf area compared to associated shrubs; about twice as large as the leaf areas of associated greenleaf manzanita (Arctostaphylos patula) and redstem ceanothus (Ceanothus sanguineus) [40]. Agee and Lolley [6] placed oceanspray in fuel type 2: shrubs with thick stems but thin leaves.

Fire regimes: Plant communities with oceanspray display a wide range of fire regimes. Oceanspray is most often dominant in dry Douglas-fir forests but is also common in dry ponderosa pine (Pinus ponderosa) and mesic western redcedar-western hemlock forests. Historical fire regimes of these coniferous forests are a continuum from mostly frequent, low-severity surface fires for ponderosa pine; to mixed surface and crown fires for Douglas-fir; to mostly long return-interval, crown fires for western redcedar-western hemlock types. Oceanspray also occurs in less widely distributed conifer and nonconifer types that historically experienced a similarly wide range of fire regimes. These types include Port-Orford-cedar, redwood, and knobcone pine coniferous communities and Oregon white oak and chaparral communities. See Habitat Types and Plant Communities for details on communities with oceanspray.

Douglas-fir: Douglas-fir forests with oceanspray experience low- to moderate-severity surface, stand-replacement surface, and crown fires (review by [204]). Because Douglas-fir forests with a large component of oceanspray occur mainly at low elevations, human-ignited fires can be frequent [35]. A study in the Cascade Range of Oregon found the ages of coast Douglas-fir/oceanspray stands that established after major fires (those killing ≥50% of the overstory) ranged from 75 to 450 years. Mean return intervals of major fires ranged from 72 years for the coast Douglas-fir/oceanspray/Indian's-dream (Aspidotis densa) phase of the coast Douglas-fir/oceanspray/grass type to 111 years for the coast Douglas-fir/oceanspray/vine maple type. The coast Douglas-fir/oceanspray/Indian's-dream phase is the driest phase of the coast Douglas-fir/oceanspray/grass type; the coast Douglas-fir/oceanspray/vine maple type occurs on ridges that may burn less frequently than other coast Douglas-fir/oceanspray types [153].

Chappell and Giglio [32] found fire was the major natural disturbance in coast Douglas-fir-Pacific madrone/oceanspray associations near the Puget Trough of Washington. Their study of over 50 coast Douglas-fir-Pacific madrone sites across west-central Washington found all sites had experienced fire at least once in 130 years [32]. Agee [4] reported a fire regime of mostly mixed moderate-severity surface and crown fires—with some low-severity surface fires—for coast Douglas-fir-Pacific madrone forests in the area.

Weisburg [234] found coast Douglas-fir forests in the west-central portion of the Cascade Range of Oregon showed a mean fire-return interval of 97 years from 1475 to 1996; the mean interval was 197 years when low-severity fires were excluded. Low- to moderate-severity fires averaged 58% of all fires. Basal area of understory shrubs, which included oceanspray, decreased with time since fire (R²=0.51) [234].

Ponderosa pine: Ponderosa pine communities historically experienced mostly low-severity surface fires (for example, [16,82]). Pacific ponderosa pine forests of the Blue Mountains of Washington and Oregon had mostly low-severity surface fires averaging every 6 to 22 years (review by [244]). This fire regime favored oceanspray; Wright [244] characterized oceanspray and other shrubs in these communities as "vigorous sprouters and very tolerant of fire". Very frequent fire-return intervals (<5 years), however, resulted in greater cover of bluebunch wheatgrass and other bunchgrasses than cover of shrubs [244]. Pacific ponderosa pine forests of northern Idaho and western Montana historically experienced similar fire behavior and fire-return intervals. Rocky Mountain Douglas-fir generally becomes dominant in these forests with fire exclusion (review by [244]).

Western redcedar-western hemlock: Historical fire regimes in this type were variable; some stands experienced fire only rarely but others burned at moderate intervals. Stand-replacement fires are more common in this type than understory fires. Seral forests—where oceanspray is most likely to occur—historically burned about every 50 to 100 years, usually with surface or mixed surface and crown fires. Surface fires in seral western redcedar-western hemlock forests may affect the composition of understory shrubs such as oceanspray but typically have little effect on overstory composition. Western redcedar-western hemlock forests of northern Idaho, in which oceanspray is prevalent, are typically cooler and damper than most conifer types, with a sparse understory, deep duff, and moist large woody fuels. The fire-return interval ranges from 25 to >500 years for these forests (review by [201]).

Port-Orford-cedar: Port-Orford-cedar forests experience mixed-severity and stand-replacement fires at short to long intervals. Port-Orford-cedar is somewhat unusual in that it persists from early-seral to climax stages of postfire succession [257]. Oceanspray is unlikely to persist in closed-canopy, old-growth Port-Orford-cedar (see Successional Status), but may persist in seral Port-Orford-cedar communities.

Redwood: These forests historically experienced short to long fire-return intervals [76,119], with surface fires at short to moderate intervals (10-100 years) most common [119]. Oceanspray is unlikely to persist in closed-canopy, old-growth redwood forests (see Successional Status) but may persist in seral communities.

Mixed-conifer forests of Oregon and California: These mixed-conifer forests historically had a regime of mostly low-severity surface fires on the hot, dry sites [34] where oceanspray is most likely to occur (see Site Characteristics). Fire-return intervals varied from about 10 to 80 years (review by [34]). A study in the Klamath Mountains of California found that fires in mixed-conifer forests with oceanspray were historically mostly small and of mixed severity. Fire activity declined in the 1950s, with the advent of fire exclusion [239]. Sierra Nevada lodgepole pine-white fir communities in Emigrant Basin Wilderness Area, California, historically had fire-return intervals averaging 57 years [77].

Oregon white oak: Oregon white oak communities historically had short- to moderate-interval surface fires. White and others [238] report that Oregon white oak in Oregon's Klamath Mountains experienced fire every 3 to 20 years. A fire history study of Oregon white oak-coast Douglas-fir mosaics of Vancouver Island, British Columbia, found that prior to European settlement, the island experienced mostly low-severity surface fires. Native Americans apparently burned these mosaics several times/decade, so fire-caused mortality of overstory trees was negligible. These intervals were much shorter [147] than the 50- to 100-year intervals Agee [4] estimated for coast Douglas-fir forests on the adjacent mainland.

Knobcone pine and chaparral: Some communities where oceanspray is important experience frequent crown fires. Knobcone pine communities of southern Oregon and coastal California require frequent (<60-year intervals) [230], stand-replacement fires to maintain knobcone pine as the community dominant [39,230]. Likewise, chamise communities of southern Oregon and California require stand-replacement fires at <60-year intervals to maintain chamise as the community dominant [188].

Other shrublands: In Redwood National Park the bald-hill shrubland communities, of which oceanspray is an important component, were historically maintained by very frequent fire. Native Americans may have burned the bald hills at least every other year. Fire scars on Oregon white oaks adjacent to the hills show fire-return intervals of about 2 years; 10 years was the maximum time between fires that left scars. Coast Douglas-fir seedlings are encroaching on the hills with fire exclusion; the former shrubby balds will likely succeed to redwood/Douglas-fir forests without the return of very frequent fire [211].

See the Fire Regime Table for further information on fire regimes of vegetation communities in which oceanspray 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".

MANAGEMENT CONSIDERATIONS

• FEDERAL LEGAL STATUS
• OTHER STATUS
• IMPORTANCE TO WILDLIFE AND LIVESTOCK
• VALUE FOR REHABILITATION OF DISTURBED SITES
• OTHER USES
• OTHER MANAGEMENT CONSIDERATIONS

However, because this species is common and readily available to wildlife and livestock on low-elevation rangelands, ungulates may make light but frequent use of oceanspray in summer [161,226]. Cattle use it as summer forage in northern Idaho [35,217] and northeastern Oregon [115].

Wildlife [125,161] and livestock [125] sometimes browse oceanspray more heavily [36,62,115], especially in late fall and winter when green forage is less available [125,197]. Snowshoe hares in the Flathead region of western Montana use the leaves and twigs for fall forage [3]. Studies on the Bitterroot National Forest and in the Rattlesnake Creek drainage of western Montana found elk, mule deer, and white-tailed deer preferred oceanspray as winter forage [109,146]. Columbian black-tailed deer in western Oregon browse oceanspray twigs in winter [47]; mule deer on the Los Padres National Forest of southern California also use oceanspray [184]. Oceanspray is heavily utilized by migrating mule deer and elk in central Washington [187].

Green clippings of oceanspray were found in dusky-footed woodrat shelters in Oregon [29], and the shrub is apparently palatable to native slugs in western Washington [31].

Palatability and/or nutritional value: Oceanspray is usually unpalatable to ungulates [157,158] and other browsing animals. A review rated its palatability as poor to fair for cattle and fair for domestic sheep [80]. A study on the Tillamook Burn of northwestern Oregon found mountain beavers browsed oceanspray less than expected based on availability [46]. New postfire sprouts are most palatable ([18,166], review by [196]). On burned sites in northern Idaho, big game species in northern Idaho preferred browsing sprouts of oceanspray and other shrubs to browsing current-year growth of shrubs on adjacent unburned sites, especially the first growing season after fire [18]. On one site, elk utilization of oceanspray increased from 1.3% before fire to 36.3% a year after prescribed fire; elk use dropped to 6.9% in postfire year 2 [127]. Asherin [18] also noted that big game species browsed oceanspray readily in postfire year 1, but use dropped after that. Browsing ungulates may pass over oceanspray sprouts if more palatable shrubs are available. On a wildfire-burned Rocky Mountain Douglas-fir/mallow ninebark habitat type on upper Selway River, northern Idaho, mule deer browsed oceanspray "minimally" in postfire years 2 and 3, while western serviceberry and Scouler willow were used heavily [105]. Following prescribed fires on the Lochsa Watershed in northern Idaho, elk preferred Scouler willow, western serviceberry, and Rocky Mountain maple sprouts to those of oceanspray [126].

Habitat: Conifer/oceanspray communities provide important habitat to a variety of wildlife species. Along the Umatilla River of Oregon, white-tailed deer used Pacific ponderosa pine-coast Douglas-fir/oceanspray and Pacific ponderosa pine/oceanspray communities more than expected based on availability (P<0.0001) [21]. On sky islands across Nevada and in western Utah, yellow-bellied marmot burrows were closely associated with oceanspray, "almost without exception" [64,65]. In the central Oregon Coast Ranges, oceanspray was found on streamside and upslope habitats where 18 of 22 small mammal species and 9 of 13 amphibian species known to the area were captured [145]. This shrub is also common in northern Idaho Pacific treefrog habitats [190].

Cover value: Oceanspray provides cover for a variety of species. Blue grouse hide beneath oceanspray and other shrubs [66]. Dense shrub understories in Rocky Mountain Douglas-fir/mallow ninebark habitat types—where oceanspray is common to codominant—provide visual and thermal cover for deer and elk; in addition, these sites supply nesting habitat, cover, and food for a variety of nongame birds and mammals [36].

VALUE FOR REHABILITATION OF DISTURBED SITES:
Oceanspray could potentially be used on Burned Area Recovery sites, although to date (2010), there was no documentation of its suvivorship after transplanting onto burns. It is used successfully for erosion control ([154], review by [196]), highway plantings, windbreaks, riparian plantings, and wildlife plantings (reviews by [196,197]). It establishes readily through natural regeneration on burned sites [35,246] (see Successional Status and Plant response to fire). In northern Idaho, for example, oceanspray dominated (48% of total understory cover) a Pacific ponderosa pine-Rocky Mountain Douglas-fir stand in early postfire succession; its size (10-15 feet (3-4.6 m)) and relative unpalatability allowed it to compete successfully with other shrubs for light, moisture, and space. Because it is a "poor forage species", researchers predicted it would dominate the burn until crowded out by conifers [203].

Oceanspray is propagated from cuttings or seed [12,113], with cuttings the usual method. A 2004 review found oceanspray seeds were "rare and costly" [196], and as of 2008, there were no published guidelines for growing this species from seed [197]. See these sources: ([74], reviews by [196,197]) for information on propagating oceanspray. Plants are available commercially [74].

Traditional uses: Native Americans used oceanspray for making implements, as medicine [255], and sometimes as food. The long, straight, hard branchwood was highly prized for making arrow shafts [50,224], as well as digging sticks, fishing hooks, and needles [73,224]. Native Americans used oceanspray for treating viral and skin diseases ([73], review by [164]) and as a tonic [73]. The bark and leaves were dried and pulverized for application to burns or sores [86]. The Pima made tea from the leaves [118], and Native Americans in the Inland Northwest ate the seeds [55].

Grazing: Oceanspray may decline with grazing despite its relative unpalatibility; its growth response in browsing and clipping studies has been mixed. In northern Idaho sites with elk, moose, mule deer, and white-tailed deer, oceanspray was more common inside than outside exclosures [8]. Another northern Idaho study in a Douglas-fir habitat type found oceanspray decreased in cattle-grazed stands [253]. Similarly, in the Bitterroot Mountains of northern Idaho, oceanspray showed greater density, cover, and frequency on ungrazed plots than on plots grazed by cattle (989 vs. 522 plants/ha; 2.6% vs. 0.6%; 4.4% vs. 1.3%, respectively, for desnity, cover, and frequency) [252]. Garrison [71] recommends ≤50% to 60% utilization of oceanspray to prevent the species' decline.

Daubenmire and Daubenmire [51] reported that in eastern Washington and northern Idaho, overgrazing of Pacific ponderosa pine/mallow ninebark stands, in which oceanspray often codominates, may result in a disclimax ponderosa pine/bluegrass (Poa spp.) community.

APPENDIX: FIRE REGIME TABLE

REFERENCES

1. Ackerly, David. 2004. Functional strategies of chaparral shrubs in relation to seasonal water deficit and disturbance. Ecological Monographs. 74(1): 25-44. [47395]

2. Adams, David L.; Mahoney, Ronald L. 1991. Effects of shade and competing vegetation on growth of western redcedar regeneration. Western Journal of Applied Forestry. 6(1): 21-22. [15151]

3. Adams, Lowell. 1959. An analysis of a population of snowshoe hares in northwestern Montana. Ecological Monographs. 29(2): 148-153. [25154]

4. Agee, James K. 1993. Fire ecology of Pacific Northwest forests. Washington, DC: Island Press. 493 p. [22247]

5. Agee, James K.; Finney, Mark; de Gouvenain, Roland. 1990. Forest fire history of Desolation Peak, Washington. Canadian Journal of Forest Research. 20: 350-356. [11035]

6. Agee, James K.; Lolley, M. Reese. 2009. Fuels and fire behavior. In: Agee, James K.; Lehmkuhl, John F., compilers. Dry forests of the northeastern Cascades fire and fire surrogate project sites, Mission Creek, Okanogan-Wenatchee National Forest. Research Paper PNW-RP-577. Portland , OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station: 35-46. [79562]

7. Agee, James K.; Wright, Clinton S.; Williamson, Nathan; Huff, Mark H. 2002. Foliar moisture content of Pacific Northwest vegetation and its relation to wildland fire behavior. Forest and Ecology Management. 167: 57-66. [41775]

8. Alldredge, Matthew W.; Peek, James M.; Wall, William A. 2001. Alterations of shrub communities in relation to herbivory in northern Idaho. Northwest Science. 75(2): 137-144. [53493]

9. Allen, Barbara H.; Holzman, Barbara A.; Evett, Rand R. 1991. A classification system for California's hardwood rangelands. Hilgardia. 59(2): 1-45. [17371]

10. Anderson, H. G. 1969. Growth form and distribution of vine maple (Acer circinatum) on Mary's Peak, western Oregon. Ecology. 50(1): 127-130. [8425]

11. Anderson, Howard George. 1967. The phytosociology of some vine maple communities in the Mary's Peak watershed. Corvallis, OR: Oregon State University. 118 p. Thesis. [9877]

12. Antieau, Clayton J. 1987. Holodiscus discolor. American Nurseryman. 166(2): 110. [76325]

13. Antieau, Clayton Joseph. 1985. Patterns of natural variation in oceanspray, Holodiscus discolor (Pursh) Maxim. (Rosaceae). Seattle, WA: University of Washington. 164 p. Thesis. [76401]

14. Antos, J. A.; Habeck, J. R. 1981. Successional development in Abies grandis (Dougl.) Forbes forests in the Swan Valley, western Montana. Northwest Science. 55(1): 26-39. [12445]

15. Armour, Charles D.; Bunting, Stephen C.; Neuenschwander, Leon F. [n.d.]. The effect of fire intensity on understory vegetational development. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 17 p. [30855]

16. Arno, Stephen F. 1988. Fire ecology and its management implications in ponderosa pine forests. In: Baumgartner, David M.; Lotan, James E., compilers. Ponderosa pine: The species and its management: Symposium proceedings; 1987 September 29 - October 1; Spokane, WA. Pullman, WA: Washington State University, Cooperative Extension: 133-139. [9410]

17. Arno, Stephen F.; Wilson, Andrew E. 1986. Dating past fires in curlleaf mountain-mahogany communities. Journal of Range Management. 39(3): 241-243. [350]

18. Asherin, Duane A. 1975. Changes in elk use and available browse production on north Idaho winter ranges following prescribed burning. In: Hieb, Susuan R., ed. Proceedings, elk logging-roads symposium; 1975 December 16-17; Moscow, ID. Moscow, ID: University of Idaho: 122-134. [17049]

19. Atzet, Thomas; White, Diane E.; McCrimmon, Lisa A.; Martinez, Patricia A.; Fong, Paula Reid; Randall, Vince D., tech. coords. 1996. Field guide to the forested plant associations of southwestern Oregon. Tech. Pap. R6-NR-ECOL-TP-17-96. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. Available online: https://www.fs.usda.gov /r6/rogue-siskiyou/publications/plant-associations.shtml [2008, September 12]. [49881]

20. Bailey, Arthur Wesley. 1966. Forest associations and secondary succession in the southern Oregon Coast Range. Corvallis, OR: Oregon State University. 166 p. Thesis. [5786]

21. Bell, Jack H.; Lauer, Jerry L.; Peek, James M. 1992. Habitat use patterns of white-tailed deer, Umatilla River, Oregon. Northwest Science. 66(3): 160-171. [19276]

22. Berch, Shannon M.; Gamiet, Sharmin; Deom, Elisabeth. 1988. Mycorrhizal status of some plants of southwestern British Columbia. Canadian Journal of Botany. 66: 1924-1928. [8841]

23. Blower, Dan. 1982. Key winter forage plants for B.C. ungulates. Victoria, BC: British Columbia Ministry of the Environment, Terrestrial Studies Branch. 57 p. [17065]

24. Bolsinger, Charles L. 1989. California's western juniper and pinyon-juniper woodlands: area, stand characteristics, wood volume, and fenceposts. Res. Bull. PNW-RB-166. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 37 p. [10365]

25. Bradley, Anne F.; Fischer, William C.; Noste, Nonan V. 1992. Fire ecology of the forest habitat types of eastern Idaho and western Wyoming. Gen. Tech. Rep. INT-290. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 92 p. [19557]

26. Bradshaw, Larry S. [n.d.]. Post-fire vegetation and fuel succession in the White Cap Wilderness Study Area: 1972-1980. Final Report: Cooperative Agreement 22-C-3-INT-28-CA. On file with: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 23 p. [20955]

27. Brown, J. K. 1976. Estimating shrub biomass from basal stem diameters. Canadian Journal of Forest Research. 6: 153-358. [10107]

28. Brown, James K.; Smith, Jane Kapler, eds. 2000. 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. 257 p. [36581]

29. 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. [18163]

30. Carey, Andrew B.; Kershner, Janet; Biswell, Brian; Dominguez de Toledo, Laura. 1999. Ecological scale and forest development: squirrels, dietary fungi, and vascular plants in managed and unmanaged forests. Wildlife Monographs No. 142. Washington, DC: The Wildlife Society. 71 p. [30476]

31. Cates, Rex G.; Orians, Gordon H. 1975. Successional status and the palatability of plants to generalized herbivores. Ecology. 56: 410-418. [15989]

32. 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. [40472]

33. Chappell, Christopher B.; Kagan, Jimmy. 2001. 2. Westside oak and dry Douglas-fir forest and woodland. In: Chappell, Christopher B.; Crawford, Rex C.; Barrett, Charley; Kagan, Jimmy; Johnson, David H.; O'Mealy, Mikell; Green, Greg A.; Ferguson, Howard L.; Edge, W. Daniel; Greda, Eva L.; O'Neil, Thomas A. Wildlife habitats: descriptions, status, trends, and system dynamics. In: Johnson, David H.; O'Neil, Thomas A., eds. Wildlife-habitat relationships in Oregon and Washington. Corvallis, OR: Oregon State University Press: 26-28. [68065]

34. Chappell, Christopher B.; Kagan, Jimmy. 2001. 3. Southwest Oregon mixed conifer-hardwood forest. In: Chappell, Christopher B.; Crawford, Rex C.; Barrett, Charley; Kagan, Jimmy; Johnson, David H.; O'Mealy, Mikell; Green, Greg A.; Ferguson, Howard L.; Edge, W. Daniel; Greda, Eva L.; O'Neil, Thomas A. Wildlife habitats: descriptions, status, trends, and system dynamics. In: Johnson, David H.; O'Neil, Thomas A., eds. Wildlife-habitat relationships in Oregon and Washington. Corvallis, OR: Oregon State University Press: 28-30. [68066]

35. Cholewa, Anita F. 1977. Successional relationships of vegetational composition to logging, burning, and grazing in the Douglas-fir/Physocarpus habitat type of northern Idaho. Moscow, ID: University of Idaho. 65 p. [+ appendices]. Thesis. [29853]

36. Cholewa, Anita F.; Johnson, Frederic D. 1983. Secondary succession in the Pseudotsuga menziesii/Physocarpus malvaceus association. Northwest Science. 57(4): 273-282. [11402]

37. Christianson, Steven P.; Adams, David L.; Grahm, Russell T. 1984. First season survival and growth of Douglas-fir planted in north Idaho shrubfields. Tech. Rep. 16. Moscow, ID: University of Idaho, Forest, Wildlife and Range Experiment Station. 6 p. [7256]

38. Clarke, Sharon E.; Garner, Mark W.; McIntosh, Bruce A.; Sedell, James R. 1997. Section 3--Landscape-level ecoregions for seven contiguous watersheds, northeast Oregon and southeast Washington. In: Clarke, Sharon E.; Bryce, Sandra A., eds. Hierarchical subdivisions of the Columbia Plateau and Blue Mountains ecoregions, Oregon and Washington. Gen. Tech. Rep. PNW-GTR-395. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station: 56-113. [28539]

39. Colwell, Wilmer L., Jr. 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. [50059]

40. Conard, Susan G.; Sparks, Steven R.; Regelbrugge, Jon C. 1997. Comparative plant water relations and soil water depletion patterns of three seral shrub species on forest sites in southwestern Oregon. Forest Science. 43(3): 336-347. [76322]

41. Conrad, C. Eugene. 1987. Common shrubs of chaparral and associated ecosystems of southern California. Gen. Tech. Rep. PSW-99. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station. 86 p. [4209]

42. Crane, M. F.; Fischer, William C. 1986. Fire ecology of the forest habitat types of central Idaho. Gen. Tech. Rep. INT-218. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 85 p. [5297]

43. Crawford, Rex C. 2001. 6. Lodgepole pine forest and woodlands. In: Chappell, Christopher B.; Crawford, Rex C.; Barrett, Charley; Kagan, Jimmy; Johnson, David H.; O'Mealy, Mikell; Green, Greg A.; Ferguson, Howard L.; Edge, W. Daniel; Greda, Eva L.; O'Neil, Thomas A. Wildlife habitats: descriptions, status, trends, and system dynamics. In: Johnson, David H.; O'Neil, Thomas A., eds. Wildlife-habitat relationships in Oregon and Washington. Corvallis, OR: Oregon State University Press: 34-35. [68069]

44. Crawford, Rex C.; Kagan, Jimmy. 2001. 14. Eastside canyon shrublands. In: Chappell, Christopher B.; Crawford, Rex C.; Barrett, Charley; Kagan, Jimmy; Johnson, David H.; O'Mealy, Mikell; Green, Greg A.; Ferguson, Howard L.; Edge, W. Daniel; Greda, Eva L.; O'Neil, Thomas A. Wildlife habitats: descriptions, status, trends, and system dynamics. In: Johnson, David H.; O'Neil, Thomas A., eds. Wildlife-habitat relationships in Oregon and Washington. Corvallis, OR: Oregon State University Press:46-47. [68103]

45. Cronquist, Arthur; Holmgren, Noel H.; Holmgren, Patricia K. 1997. Intermountain flora: Vascular plants of the Intermountain West, U.S.A. Vol. 3, Part A: Subclass Rosidae (except Fabales). New York: The New York Botanical Garden. 446 p. [28652]

46. Crouch, Glenn L. 1968. Clipping of woody plants by mountain beaver. Journal of Mammalogy. 49(1): 151-152. [64410]

47. Crouch, Glenn L. 1968. Forage availability in relation to browsing of Douglas-fir seedlings by black-tailed deer. Journal of Wildlife Management. 32(3): 542-553. [16105]

48. Cutright, Paul Russell. 1989. Lewis and Clark: pioneering naturalists. First Bison Book. Lincoln, NE: University of Nebraska Press. 506 p. [20300]

49. Dasgupta, Swarvanu; Qu, John J.; Hao, Xianjun; Bhoi, Sanjeeb. 2007. Evaluating remotely sensed live fuel moisture estimations for fire behavior predictions in Georgia, USA. Remote Sensing of Environment. 108(2): 138-150. [79564]

50. Daubenmire, R. 1970. Steppe vegetation of Washington. Technical Bulletin 62. Pullman, WA: Washington State University, College of Agriculture; Washington Agricultural Experiment Station. 131 p. [733]

51. Daubenmire, Rexford F.; Daubenmire, Jean B. 1968. Forest vegetation of eastern Washington and northern Idaho. Technical Bulletin 60. Pullman, WA: Washington State University, College of Agriculture; Washington Agricultural Experiment Station. 104 p. [749]

52. Dayton, William A. 1931. Important western browse plants. Misc. Publ. No. 101. Washington, DC: U.S. Department of Agriculture. 214 p. [768]

53. del Moral, Roger; Cates, Rex G. 1971. Allelopathic potential of the dominant vegetation of western Washington. Ecology. 52(6): 1030-1037. [4794]

54. Donnelly, Steve. 1993. Spring burning by habitat type in relation to artificial restoration. McCall, ID: U.S. Department of Agriculture, Forest Service, Intermountain Region, Payette National Forest. 19 p. [27626]

55. Drew, Larry Albert. 1967. Comparative phenology of seral shrub communities in the cedar/hemlock zone. Moscow, ID: University of Idaho. 108 p. Thesis. [9654]

56. Driscoll, Richard S. 1964. A relict area in the central Oregon juniper zone. Ecology. 45(2): 345-353. [5181]

57. Dyrness, C. T.; Franklin, J. F.; Moir, W. H. 1974. A preliminary classification of forest communities in the central portion of the western Cascades in Oregon. Bulletin No. 4. Seattle, WA: University of Washington, Ecosystem Analysis Studies, Coniferous Forest Biome. 123 p. [8480]

58. Ferguson, Robert B. 1983. Use of rosaceous shrubs for wildland plantings in the Intermountain West. In: Monsen, Stephen B.; Shaw, Nancy, comps. Managing Intermountain rangelands--improvement of range and wildlife habitats; Proceedings of symposia; 1981 September 15-17; Twin Falls, ID; 1982 June 22-24; Elko, NV. Gen. Tech. Rep. INT-157. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: 136-149. [915]

59. Finney, Mark A.; Martin, Robert E. 1989. Fire history in a Sequoia sempervirens forest at Salt Point State Park, California. Canadian Journal of Forest Research. 19: 1451-1457. [9845]

60. Fischer, William C.; Bradley, Anne F. 1987. Fire ecology of western Montana forest habitat types. Gen. Tech. Rep. INT-223. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 95 p. [633]

61. Fischer, William C.; Clayton, Bruce D. 1983. Fire ecology of Montana forest habitat types east of the Continental Divide. Gen. Tech. Rep. INT-141. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 83 p. [923]

62. Flessner, T. R.; Darris, D. C.; Lambert, S. M. 1992. Seed source evaluation of four native riparian shrubs for streambank rehabilitation in the Pacific Northwest. In: Clary, Warren P.; McArthur, E. Durant; Bedunah, Don; Wambolt, Carl L., compilers. Proceedings--symposium on ecology and management of riparian shrub communities; 1991 May 29-31; Sun Valley, ID. Gen. Tech. Rep. INT-289. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 155-162. [19111]

63. Flora of North America Association. 2010. Flora of North America: The flora, [Online]. Flora of North America Association (Producer). Available: http://www.fna.org/FNA. [36990]

64. Floyd, Chris H. 2004. Marmot distribution and habitat associations in the Great Basin. Western North American Naturalist. 64(4): 471-481. [76354]

65. Floyd, Christian Hollace. 2003. Ecological genetics of dispersal and mating system in populations of yellow-bellied marmots (Marmota flaviventris). Davis, CA: University of California, Davis. 148 p. Dissertation. [76400]

66. Fowle, C. David. 1960. A study of the blue grouse (Dendragapus obscurus Say) on Vancouver Island, British Columbia. Canadian Journal of Zoology. 38(4): 701-713. [34529]

67. 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. [8207]

68. Franklin, Jerry F. 1988. Pacific Northwest forests. In: Barbour, Michael G.; Billings, William Dwight, eds. North American terrestrial vegetation. New York: Cambridge University Press: 103-130. [13879]

69. Freedman, June D.; Habeck, James R. 1985. Fire, logging, and white-tailed deer interrelationships in the Swan Valley, northwestern Montana. In: Lotan, James E.; Brown, James K., compilers. Fire's effects on wildlife habitat--symposium proceedings; 1984 March 21; Missoula, MT. Gen. Tech. Rep. INT-186. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 23-35. [8319]

70. Garrison, George A. 1953. Effects of clipping on some range shrubs. Journal of Range Management. 6(5): 309-317. [995]

71. Garrison, George A. 1972. Carbohydrate reserves and response to use. In: McKell, Cyrus M.; Blaisdell, James P.; Goodin, Joe R., eds. Wildland shrubs--their biology and utilization: Proceedings of a symposium; 1971 July; Logan, UT. Gen. Tech. Rep. INT-1. Ogden, UT; U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: 271-278. [997]

72. Goheen, Donald; Frankel, Susan. 1993. Using diverse plant species to maintain forest health. In: Landis, Thomas D., tech. coord. Proceedings, Western Forest Nursery Association; 1992 September 14-18; Fallen Leaf Lake, CA. Gen. Tech. Rep. RM-221. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 18-20. [22068]

73. Gonzalez-Laredo, Rubin F.; Chaidez-Gonzalez, Judith; Ahmed, Ahmed A.; Karchesy, Joseph J. 1997. A stilbene xyloside from Holodiscus discolor bark. Phytochemistry. 46(1): 175-176. [76317]

74. Gonzalves, Pete; Darris, Dale. 2007. Plant fact sheet--Oceanspray: Holodiscus discolor (Pursh.) Maxim, [Online]. In: Plant profile--Holodiscus discolor. In: PLANTS database. Baton Rouge, LA: U.S. Department of Agriculture, Natural Resources Conservation Service, National Plant Data Center (Producer). Available: https://plants.usda.gov /factsheet/pdf/fs_hodi.pdf. In: https://plants.usda.gov / [2010, April 28]. [79499]

75. Green, R. N.; Courtin, P. J.; Klinka, K.; Slaco, R. J.; Ray, C. A. 1984. Site diagnosis, tree species selection, and slashburning guidelines for the Vancouver Forest Region. Land Management Handbook Number 8 [Abridged version]. Burnaby, BC: Ministry of Forests, Vancouver Forest Region. 143 p. [9475]

76. Greenlee, Jason M.; Langenheim, Jean H. 1990. Historic fire regimes and their relation to vegetation patterns in the Monterey Bay area of California. The American Midland Naturalist. 124(2): 239-253. [15144]

77. Greenlee, John M. 1973. A study of the fire ecology of the Emigrant Basin Primitive Area: Stanislaus National Forest. [Project No. 14]. Sonora, CA: U.S. Department of Agriculture, Forest Service, Stanislaus National Forest, Summit Ranger District, Pinecrest Ranger Station. 64 p. [21257]

78. Griffin, James R. 1974. Notes on environment, vegetation and flora: Hastings Natural History Reservation. Carmel Valley, CA: University of California, Hastings Natural History Reservation. Unpublished paper on file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 90 p. [10531]

79. Gruell, George E. 2001. Fire in Sierra Nevada forests: A photographic interpretation of ecological change since 1849. Missoula, MT: Mountain Press Publishing Company. 238 p. [43843]

80. Gullion, Gordon W. 1964. Wildlife uses of Nevada plants. Contributions toward a flora of Nevada: No. 49. CR-24-64. Beltsville, MD: U.S. Department of Agriculture, Agricultural Research Service, Crops Research Division; Washington, DC: U.S. National Arboretum, Herbarium. 170 p. [6729]

81. Habeck, James R. 1968. Forest succession in the Glacier Park cedar-hemlock forests. Ecology. 49(5): 872-880. [6479]

82. Habeck, James R. 1990. Old-growth ponderosa pine-western larch forests in western Montana: ecology and management. Northwest Environmental Journal. 6(2): 271-292. [17661]

83. Haire, Sandra L.; McGarigal, Kevin. 2008. Inhabitants of landscape scars: succession of woody plants after large, severe forest fires in Arizona and New Mexico. The Southwestern Naturalist. 53(2): 146-161. [70959]

84. Hall, Frederick C. 1973. Plant communities of the Blue Mountains in eastern Oregon and southeastern Washington. R6 Area Guide 3-1. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 82 p. [1059]

85. Halofsky, Jessica E.; Hibbs, David E. 2009. Controls on early post-fire woody plant colonization in riparian areas. Forest Ecology and Management. 258(7): 1350-1358. [77249]

86. Halverson, Nancy M., comp. 1986. Major indicator shrubs and herbs on national forests of western Oregon and southwestern Washington. R6-TM-229. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 180 p. [3233]

87. Halverson, Nancy M.; Topik, Christopher; Van Vickle, Robert. 1986. Plant association and management guide for the western hemlock zone: Mt. Hood National Forest. R6-ECOL-232A. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 111 p. [1068]

88. Hann, Wendel; Havlina, Doug; Shlisky, Ayn; [and others]. 2008. Interagency fire regime condition class guidebook. Version 1.3, [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). 119 p. Available: http://frames.nbii.gov/frcc/documents/FRCC_Guidebook_2008.07.10.pdf [2010, 3 May]. [70966]

89. Harrington, Constance A.; McGrath, James M.; Kraft, Joseph M. 1999. Propagating native species: experience at the Wind River Nursery. Western Journal of Applied Forestry. 14(2): 61-64. [30058]

90. Harris, Gregory Dean. 2004. Integrating a shrub growth model with remote sensing and geographic information system data to predict shrub spatial growth patterns in a post fire environment. Pullman, WA: Washington State University. 130 p. Dissertation. [76399]

91. Harrod, Richy J.; Fonda, Richard W.; McGrath, Mara K. 2007. Role of fire in restoration of a ponderosa pine forest, Washington. In: Butler, Bret W.; Cook, Wayne, comps. The fire environment--innovations, management, and policy; conference proceedings; 2007 March 26-30; Destin, FL. Proceedings RMRS-P-46CD. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 315-328. CD-ROM. [71109]

92. Harrod, Richy J.; Knecht, Dottie E.; Kuhlmann, Ellen E.; Ellis, Mark W.; Davenport, Roberta. 1997. Effects of the Rat and Hatchery Creek fires on four rare plant species. In: Greenlee, Jason M., ed. Proceedings, 1st conference on fire effects on rare and endangered species and habitats; 1995 November 13-16; Coeur d'Alene, ID. Fairfield, WA: International Association of Wildland Fire: 311-319. [28155]

93. Hebda, R. J.; Chinnappa, C. C.; Smith, B. M. 1988. Pollen morphology of the Rosaceae of western Canada. II. Dryas, Fragaria, Holodiscus. Canadian Journal of Botany. 66: 595-612. [4514]

94. Hemstrom, Miles A.; Logan, Sheila E.; Pavlat, Warren. 1987. Plant association and management guide: Willamette National Forest. R6-Ecol 257-B-86. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 312 p. [13402]

95. Henderson, Jan A.; Peter, David H.; Lesher, Robin D.; Shaw, David C. 1989. Forested plant associations of the Olympic National Forest. R6-ECOL-TP 001-88. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 502 p. [23405]

96. Hickey, William O. 1971. Response of Ceanothus sanguineus to cutting and burning at various stages of phenology. Moscow, ID: University of Idaho. 39 p. Thesis. [9777]

97. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992]

98. Hines, William Wester. 1971. Plant communities in the old-growth forests of north coastal Oregon. Corvallis, OR: Oregon State University. 146 p. Thesis. [10399]

99. Hitchcock, C. Leo; Cronquist, Arthur. 1973. Flora of the Pacific Northwest. Seattle, WA: University of Washington Press. 730 p. [1168]

100. Hitchcock, C. Leo; Cronquist, Arthur; Ownbey, Marion; Thompson, J. W. 1961. Vascular plants of the Pacific Northwest. Part 3: Saxifragaceae to Ericaceae. Seattle, WA: University of Washington Press. 614 p. [1167]

101. Irwin, Larry L.; Peek, James M. 1979. Shrub production and biomass trends following five logging treatments within the cedar-hemlock zone of northern Idaho. Forest Science. 25(3): 415-426. [16511]

102. Jantova, S.; Nagy, M.; Ruzekova, L.; Grancai, D. 2000. Antibacterial activity of plant extracts from the families Fabaceae, Oleaceae, Philadelphaceae, Rosaceae and Staphyleaceae. Phytotherapy Research. 14(8): 601-603. [76312]

103. 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. [36715]

104. Kartesz, John Thomas. 1988. A flora of Nevada. Reno, NV: University of Nevada. 1729 p. [In 2 volumes]. Dissertation. [42426]

105. Keay, Jeffrey A. 1977. Relationship of habitat use patterns and forage preferences of white-tailed and mule deer to post-fire vegetation, upper Selway River. Moscow, ID: University of Idaho. 76 p. Thesis. [1316]

106. Keeler-Wolf, Todd. 1986. An ecological survey of the proposed Stone Corral - Josephine Peridotite Research Natural Area (L. E. Horton - Darlingtonia Bog Research Natural Area) on the Six Rivers National Forest, Del Norte County, California. Purchase order # 40-9AD6-5-907. Unpublished report on file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 69 p. [12307]

107. Kelpsas, B. R. 1978. Comparative effects of chemical, fire, and machine site preparation in an Oregon coastal brushfield. Corvallis, OR: Oregon State University. 97 p. Thesis. [6986]

108. Kessell, Stephen R. 1979. Comparison of community stratification methods in Mount Rainier National Park and Glacier National Park. Unpublished preliminary report on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Lab, Missoula, MT. 154 p. [6678]

109. Klebenow, Donald A. 1965. A montane forest winter deer habitat in western Montana. Journal of Wildlife Management. 29(1): 27-33. [8430]

110. Klinka, K.; Green, R. N.; Courtin, P. J.; Nuszdorfer, F. C. 1984. Site diagnosis, tree species selection, and slashburning guidelines for the Vancouver Forest Region, British Columbia. Land Management Report No. 25. Victoria, BC: Ministry of Forests, Information Services Branch. 180 p. [15448]

111. 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. [10703]

112. Kovalchik, Bernard L.; Clausnitzer, Rodrick R. 2004. Classification and management of aquatic, riparian, and wetland sites on the national forests of eastern Washington: series description. Gen. Tech. Rep. PNW-GTR-593. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 354 p. [53329]

113. Kruckeberg, A. R. 1982. Gardening with native plants of the Pacific Northwest. Seattle, WA: University of Washington Press. 252 p. [9980]

114. Kruckeberg, Arthur R. 1977. Manzanita (Arctostaphylos) hybrids in the Pacific Northwest: effects of human and natural disturbance. Systematic Botany. 2(4): 233-250. [13561]

115. Krueger, W. C.; Vavra, M.; Wheeler, W. P. 1980. Plant succession as influenced by habitat type, grazing management, and reseeding on a northeast Oregon clearcut. In: 1980 progress report--research in rangeland management. Special Report 586. Corvallis, OR: Oregon State University, Agricultural Experiment Station: 32-37. In cooperation with: U.S. Department of Agriculture, SEA-AR. [2749]

116. Kuchler, A. W. 1964. Ponderosa shrub forest (Pinus). In: Kuchler, A. W. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society: 10. [67010]

117. Lackschewitz, Klaus. 1991. Vascular plants of west-central Montana--identification guidebook. Gen. Tech. Rep. INT-227. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 648 p. [13798]

118. Laferriere, Joseph E.; Weber, Charles W.; Kohlhepp, Edwin A. 1991. Mineral composition of some traditional Mexican teas. Plant Foods for Human Nutrition. 41(3): 277-282. [76358]

119. LANDFIRE Rapid Assessment. 2005. Potential Natural Vegetation Group (PNVG) R1SESE--redwood, [Online]. In: Rapid assessment reference condition models. In: LANDFIRE. Washington, DC: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory; U.S. Geological Survey; The Nature Conservancy (Producers). Available: https://www.landfire.gov /zip/CA/R1SESE.pdf [2010, 14 April]. [79490]

120. 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: https://www.landfire.gov /downloadfile.php?file=RA_Modeling_Manual_v2_1.pdf [2007, May 24]. [66741]

121. LANDFIRE Rapid Assessment. 2007. Rapid assessment reference condition models, [Online]. 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: https://www.landfire.gov /models_EW.php [2008, April 18] [66533]

122. Larsen, Eric M.; Morgan, John T. 1998. Management recommendations for Washington's priority habitats: Oregon white oak woodlands. Olympia, WA: Washington Department of Fish and Wildlife. 37 p. [52756]

123. Larsen, J. A. 1925. Natural reproduction after forest fires in northern Idaho. Journal of Agricultural Research. 30(12): 1177-1197. [13193]

124. Lathrop, Earl W.; Martin, Bradford D. 1982. Response of understory vegetation to prescribed burning in yellow pine forests of Cuyamaca Rancho State Park, California. Aliso. 10(2): 329-343. [15943]

125. Lee, Lyndon C.; Pfister, Robert D. 1978. A training manual for Montana forest habitat types. Missoula, MT: University of Montana, School of Forestry, Montana Forest and Conservation Experiment Station. 142 p. [1434]

126. Leege, Thomas A. 1968. Prescribed burning for elk in northern Idaho. In: Proceedings, annual Tall Timbers fire ecology conference; 1968 March 14-15; Tallahassee, FL. No 8. Tallahassee, FL: Tall Timbers Research Station: 235-253. [5287]

127. Leege, Thomas A. 1969. Burning seral brush ranges for big game in northern Idaho. Transactions, North American Wildlife and Natural Resources Conference. 34: 429-438. [144]

128. Leege, Thomas A. 1974. Changes in browse utilization after burning. Job No. 2: July 1,1966 to March 31, 1974. Study II: Spring vs. fall burning for range rehabilitation. Final report: Elk ecology. Federal Aid to Wildlife Restoration Project: W-160-R. Boise, ID: Idaho Fish and Game Department. 5 p. [42144]

129. Leege, Thomas A. 1978. Changes in browse intercept, production and seedlings after burning--Holly Creek. Job No. 2: July 1, 1965 to June 30, 1978. Study I: Range rehabilitation by spring burning. Job Completion Report: Elk ecology. Federal Aid to Wildlife Restoration Project: W-160-R. Boise, ID: Idaho Department of Fish and Game. 11 p. [17170]

130. Leege, Thomas A. 1978. Changes in browse production after burning vs. slashing and burning on the four cardinal aspects--Polar Ridge. Job No. 3: July 1, 1967 to June 30, 1978. Study I: Range rehabilitation by spring burning. Job Completion Report: Elk ecology. Federal Aid to Wildlife Restoration Project: W-160-R. Boise, ID: Idaho Department of Fish and Game. 20 p. [17171]

131. Leege, Thomas A. 1979. Effects of repeated prescribed burns on northern Idaho elk browse. Northwest Science. 53(2): 107-113. [5116]

132. Leege, Thomas A.; Hickey, William O. 1966. Lochsa elk study. Job No. 8: July 1, 1965 to June 30, 1966. Job Completion Report: Big game surveys and investigations. Federal Aid to Wildlife Restoration Project: W-85-R-17. Boise, ID: State of Idaho Fish and Game Department. 22 p. [16759]

133. Leege, Thomas A.; Hickey, William O. 1971. Sprouting of northern Idaho shrubs after prescribed burning. Journal of Wildlife Management. 35(3): 508-515. [1437]

134. Lentile, Leigh B.; Morgan, Penelope; Hudak, Andrew T.; Bobbitt, Michael J.; Lewis, Sarah A.; Smith, Alistair M. S.; Robichaud, Peter R. 2007. Post-fire burn severity and vegetation response following eight large wildfires across the western United States. Fire Ecology. 3(1): 91-108. [70228]

135. Lesher, Robin D.; Henderson, Jan A. 1989. Indicator species of the Olympic National Forest. R6-ECOL-TP003-88. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 79 p. [15376]

136. Ley, Arline. 1943. A taxonomic revision of the genus Holodiscus (Rosaceae). Bulletin of the Torrey Botanical Club. 70(3): 275-288. [142]

137. Long, James N. 1977. Trends in plant species diversity associated with development in a series of Pseudotsuga menziesii/Gaultheria shallon stands. Northwest Science. 51(2): 119-130. [10152]

138. Luoma, Daniel L.; Frenkel, Robert E.; Trappe, James M. 1991. Fruiting of hypogeous fungi in Oregon Douglas-fir forests: seasonal and habitat variation. Mycologia. 83(3): 335-353. [21391]

139. Lyon, L. Jack; Mueggler, Walter F. 1968. Herbicide treatment of north Idaho browse evaluated six years later. Journal of Wildlife Management. 32(3): 538-541. [8428]

140. Lyon, L. Jack; Stickney, Peter F. 1976. Early vegetal succession following large northern Rocky Mountain wildfires. In: Proceedings, Tall Timbers fire ecology conference and Intermountain Fire Research Council fire and land management symposium; 1974 October 8-10; Missoula, MT. No. 14. Tallahassee, FL: Tall Timbers Research Station: 355-373. [1496]

141. MacKinnon, A.; Meidinger, D.; Klinka, K. 1992. Use of the biogeoclimatic ecosystem classification system in British Columbia. Forestry Chronicle. 68(1): 100-120. [18845]

142. 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. [7608]

143. Martin, Bradford D. 1981. Vegetation responses to prescribed burning in a mixed-conifer woodland, Cuyamaca Rancho State Park, California. Loma Linda, CA: Loma Linda University. 112 p. Thesis. [64684]

144. Martin, Bradford D. 1982. Vegetation responses to prescribed burning in Cuyamaca Rancho State Park, California. In: Conrad, C. Eugene; Oechel, Walter C., technical coordinators. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems; 1981 June 22-26; San Diego, CA. Gen. Tech. Rep. PSW-58. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 617. [6088]

145. McComb, William C.; McGarigal, Kevin; Anthony, Robert G. 1993. Small mammal and amphibian abundance in streamside and upslope habitats of mature Douglas-fir stands, western Oregon. Northwest Science. 67(1): 7-15. [20564]

146. McCulloch, Clay Y., Jr. 1955. Utilization of winter browse on wilderness big game range. Journal of Wildlife Management. 19(2): 206-215. [7933]

147. McDadi, Omar; Hebda, Richard J. 2008. Change in historic fire disturbance in a Garry oak (Quercus garryana) meadow and Douglas-fir (Pseudotsuga menziesii) mosaic, University of Victoria, British Columbia, Canada: a possible link with First Nations and Europeans. Forest Ecology and Management. 256(10): 1704-1710. [72957]

148. McDonald, G. I.; Harvey, A. E.; Tonn, J. R. 2000. Fire, competition and forest pests: landscape treatment to sustain ecosystem function. In: Neuenschwander, Leon F.; Ryan, Kevin C., tech. eds. Crossing the millennium: integrating spatial technologies and ecological principles for a new age in fire management: Proceedings, Joint Fire Sciences conference and workshop; 1999 June 15-17; Boise, ID. Vol. II. Moscow, ID: University of Idaho; Boise, ID: International Association of Wildland Fire: 195-211. [41176]

149. 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. [50055]

150. McGee, Ann; Feller, M. C. 1993. Seed banks of forested and disturbed soils in southwestern British Columbia. Canadian Journal of Botany. 71: 1574-1583. [25756]

151. McKenzie, Donald; Halpern, Charles B. 1999. Modeling the distributions of shrub species in Pacific Northwest forests. Forest Ecology and Management. 114(2/3): 293-307. [38747]

152. McMinn, Robert G. 1952. The role of soil drought in the distribution of vegetation in the northern Rocky Mountains. Ecology. 33: 1-15. [1624]

153. Means, Joseph Earl. 1980. Dry coniferous forests in the western Oregon Cascades. Corvallis, OR: Oregon State University. 264 p. Dissertation. [5767]

154. 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 [2010, June 11]. [68858]

155. Miller, Margaret M.; Miller, Joseph W. 1976. Succession after wildfire in the North Cascades National Park complex. In: Proceedings, annual Tall Timbers fire ecology conference: Pacific Northwest; 1974 October 16-17; Portland, OR. No. 15. Tallahassee, FL: Tall Timbers Research Station: 71-83. [6574]

156. Minore, Don. 1978. The Dead Indian Plateau: a historical summary of forestry observations and research in a severe southwestern Oregon environment. PNW-72. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 23 p. [8423]

157. Mitchell, John E. 1983. Overstory-understory relationships: Douglas-fir forests. In: Bartlett, E. T.; Betters, David R., eds. Overstory-understory relationships in western forests. Western Regional Research Publication No. 1. Fort Collins, CO: Colorado State University, Experiment Station: 27-34. [3314]

158. Mitchell, John E.; Rodgers, Richard T. 1985. Food habits and distribution of cattle on a forest and pasture range in northern Idaho. Journal of Range Management. 38(3): 214-220. [14430]

159. Morgan, P.; Neuenschwander, L. F. [In preparation]. Seedbank contribution to shrub regeneration following clearcutting and burning. Canadian Journal of Forest Research. [29144]

160. Morgan, Penelope; Neuenschwander, Leon F. 1988. Shrub response to high and low severity burns following clearcutting in northern Idaho. Western Journal of Applied Forestry. 3(1): 5-9. [3895]

161. Morris, Melvin S.; Schmautz, Jack E.; Stickney, Peter F. 1962. Winter field key to the native shrubs of Montana. Bulletin No. 23. Missoula, MT: Montana State University, Montana Forest and Conservation Experiment Station. 70 p. [17063]

162. Morrison, Peter H.; Swanson, Frederick J. 1990. Fire history and pattern in a Cascade Range landscape. Gen. Tech. Rep. PNW-GTR-254. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 77 p. [13074]

163. Mozingo, Hugh N. 1987. Shrubs of the Great Basin: A natural history. Reno, NV: University of Nevada Press. 342 p. [1702]

164. Mrizova, M.; Lacidova, L.; Haladova, M.; Eisenrieichova, E.; Graneai, D.; Fickova, M. 2007. Cytotoxic activity of aqueous extract from Holodiscus discolor (Pursh) Maxim. leaves. Planta Medica. 73(9): 932. Abstract. [76350]

165. Mueggler, W. F. 1961. Ecology of seral shrub communities in the cedar-hemlock zone of northern Idaho. Durham, NC: Duke University. 126 p. Thesis. [9981]

166. Nelson, Jack R. 1976. Forest fire and big game in the Pacific Northwest. In: Proceedings, annual Tall Timbers fire ecology conference: Pacific Northwest; 1974 October 16-17; Portland, OR. No. 15. Tallahassee, FL: Tall Timbers Research Station: 85-102. [6464]

167. Neuenschwander, L. F. 1978. The fire induced autecology of selected shrubs of the cold desert and surrounding forests: A-state-of-the-art review. Unpublished manuscript on file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 31 p. [1747]

168. Newton, M.; Comeau, P. G. 1990. Control of competing vegetation. In: Lavender, D. P.; Parish, R.; Johnson, C. M.; Montgomery, G.; Vyse, A.; Willis, R. A.; Winston, D., eds. Regenerating British Columbia's forests. Vancouver, BC: University of British Columbia Press: 256-265. [10719]

169. Orme, Mark L.; Leege, Thomas A. 1976. Emergence and survival of redstem (Ceanothus sanguineus) following prescribed burning. In: Proceedings, Tall Timbers fire ecology conference and fire and land management symposium; 1974 October 8-10; Missoula, MT. No. 14. Tallahassee, FL: Tall Timbers Research Station: 391-420. [6273]

170. Orme, Mark L.; Leege, Thomas A. 1980. Phenology of shrubs on a north Idaho elk range. Northwest Science. 54(3): 187-198. [1800]

171. Ossinger, Mary C. 1983. The Pseudotsuga-Tsuga/Rhododendron community in the northeast Olympic Mountains. Bellingham, WA: Western Washington University. 50 p. Thesis. [11435]

172. Owens, T. E. 1982. Postburn regrowth of shrubs related to canopy mortality. Northwest Science. 56(1): 34-40. [1806]

173. Pabst, Robert J.; Spies, Thomas A. 2001. Ten years of vegetation succession on a debris-flow deposit in Oregon. Journal of the American Water Resources Association. 37(6): 1693-1708. [41709]

174. Patterson, Patricia A.; Neiman, Kenneth E.; Tonn, Jonalea. 1985. Field guide to forest plants of northern Idaho. Gen. Tech. Rep. INT-180. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 246 p. [1839]

175. Peck, JeriLynn E.; Acker, Steven A.; McKee, W. Arthur. 1995. Autecology of mosses in coniferous forests in the central western Cascades of Oregon. Northwest Science. 69(3): 184-190. [26793]

176. Pengelly, W. Leslie. 1963. Timberlands and deer in the northern Rockies. Journal of Forestry. 61: 734-740. [175]

177. Pengelly, William Leslie. 1966. Ecological effects of slash-disposal fires on the Coeur d'Alene National Forest, Idaho. Missoula, MT: U.S. Department of Agriculture, Forest Service, Northern Region. 23 p. [174]

178. Pfister, Robert D.; Kovalchik, Bernard L.; Arno, Stephen F.; Presby, Richard C. 1977. Forest habitat types of Montana. Gen. Tech. Rep. INT-34. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 174 p. [1878]

179. Pojar, J.; Klinka, K.; Meidinger, D. V. 1987. Biogeoclimatic ecosystem classification in British Columbia. Forest Ecology and Management. 22: 119-154. [7314]

180. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]

181. Rickard, W. H. 1975. Litterfall in a Douglas-fir forest near the Trojan Nuclear Power Station, Oregon. Northwest Science. 49(4): 183-189. [8178]

182. Ripple, William J. 1994. Historic spatial patterns of old forests in western Oregon. Journal of Forestry. 92(11): 45-49. [33881]

183. Roach, A. W. 1952. Phytosociology of the Nash Crater lava flows, Linn County, Oregon. Ecological Monographs. 22: 169-193. [8759]

184. Robinson, Cyril S. 1937. Plants eaten by California mule deer on the Los Padres National Forest. Journal of Forestry. 35(3): 285-292. [51853]

185. Roemer, Hans. 1993. Vegetation and ecology of Garry oak woodlands. In: Hebda, Richard J.; Aitkens, Fran, eds. Garry oak-meadow colloquium: Proceedings; 1993; Victoria, BC. Victoria, BC: Garry Oak Meadow Preservation Society: 19-24. [64527]

186. Ross, Robert L.; Hunter, Harold E. 1976. Climax vegetation of Montana: Based on soils and climate. Bozeman, MT: U.S. Department of Agriculture, Soil Conservation Service. 64 p. [2028]

187. Rummell, Robert S. 1951. Some effects of livestock grazing on ponderosa pine forest and range in central Washington. Ecology. 32(4): 594-607. [16338]

188. Rundel, Philip W.; Parsons, David J. 1979. Structural changes in chamise (Adenostoma fasciculatum) along a fire- induced age gradient. Journal of Range Management. 32(6): 462-466. [4915]

189. Sabhasri, Sanga; Ferrell, William K. 1960. Invasion of brush species into small stand openings in the Douglas-fir forests of the Willamette foothills. Northwest Science. 34(3): 77-89. [8652]

190. Schaub, David L.; Larsen, John H., Jr. 1978. The reproductive ecology of the Pacific treefrog (Hyla regilla). Herpetologica. 34(4): 409-416. [27248]

191. Schoonmaker, Peter; McKee, Arthur. 1988. Species composition and diversity during secondary succession of coniferous forests in the western Cascade Mountains of Oregon. Forest Science. 34(4): 960-979. [6214]

192. Schultz, Brad W. 1987. Ecology of curlleaf mountain mahogany (Cercocarpus ledifolius) in western and central Nevada: population structure and dynamics. Reno, NV: University of Nevada. 111 p. Thesis. [7064]

193. Scoggan, H. J. 1978. The flora of Canada. Part 3: Dicotyledoneae (Saururaceae to Violaceae). National Museum of Natural Sciences: Publications in Botany, No. 7(3). Ottawa: National Museums of Canada. 1115 p. [75493]

194. Shatford, J. P. A.; Hibbs, D. E.; Puettmann, K. J. 2007. Conifer regeneration after forest fire in the Klamath-Siskiyous: how much, how soon? Journal of Forestry. April/May: 139-146. [67813]

195. Shatford, Jeffrey P. A.; Bailey, John D.; Tappeiner, John C. 2009. Understory tree development with repeated stand density treatments in coastal Douglas-fir forests of Oregon. Western Journal of Applied Forestry. 24(1): 11-16. [73400]

196. Shaw, Nancy L. 2004. Holodiscus discolor. In: Francis, John K., ed. Wildland shrubs of the United States and its territories: thamnic descriptions: volume 1. Gen. Tech. Rep. IITF-GTR-26. San Juan, PR: U.S. Department of Agriculture, Forest Service, International Institute of Tropical Forestry; Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 379-381. [52172]

197. Shaw, Nancy L.; Hurd, Emerenciana G.; Stickney, Peter F. 2008. Holodiscus (K. Koch) Maxim.: ocean-spray. In: Bonner, Franklin T., Karrfalt, Robert P., eds. Woody plant seed manual. Agric. Handbook 727. Washington, DC: U.S. Department of Agriculture, Forest Service: 591-594. [78278]

198. Shaw, Nancy L.; Monsen, Stephen B.; Stevens, Richard. 2004. Rosaceous shrubs. In: Monsen, Stephen B.; Stevens, Richard; Shaw, Nancy L., comps. Restoring western ranges and wildlands. Gen. Tech. Rep. RMRS-GTR-136-vol-2. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 539-596. [52845]

199. Sheridan, Chris D.; Spies, Thomas A. 2005. Vegetation-environment relationships in zero-order basins in coastal Oregon. Canadian Journal of Forest Research. 35: 340-355. [60361]

200. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]

201. Shiplett, Brian; Neuenschwander, Leon F. 1994. Fire ecology in the cedar-hemlock zone of North Idaho. In: Baumgartner, David M.; Lotan, James E.; Tonn, Jonalea R., compilers. Interior cedar-hemlock-white pine forests: ecology and management: Symposium proceedings; 1993 March 2-4; Spokane, WA. Pullman, WA: Washington State University, Department of Natural Resources: 41-51. [25789]

202. Shreve, Forrest. 1927. The vegetation of a coastal mountain range. Ecology. 8(1): 27-44. [63417]

203. Simmerman, Dennis G.; Arno, Stephen F.; Harrington, Michael G.; Graham, Russell T. 1991. A comparison of dry and moist fuel underburns in ponderosa pine shelterwood units in Idaho. In: Andrews, Patricia L.; Potts, Donald F., eds. Proceedings, 11th annual conference on fire and forest meteorology; 1991 April 16-19; Missoula, MT. SAF Publication 91-04. Bethesda, MD: Society of American Foresters: 387-397. [16186]

204. Smith, Jane Kapler; Fischer, William C. 1997. Fire ecology of the forest habitat types of northern Idaho. Gen. Tech. Rep. INT-GTR-363. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 142 p. [27992]

205. Smith, W. Brad; Brand, Gary J. 1983. Allometric biomass equations for 98 species of herbs, shrubs, and small trees. Res. Note NC-299. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 8 p. [20785]

206. Stickney, Peter F. 1986. First decade plant succession following the Sundance Forest Fire, northern Idaho. Gen. Tech. Rep. INT-197. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 26 p. [2255]

207. 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, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 10 p. [20090]

208. Stickney, Peter F. 1991. Effects of fire on flora: Northern Rocky Mountain forest plants. Unpublished paper on file at: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experimental Station, Missoula, MT. 10 p. [21628]

209. Stickney, Peter F.; Campbell, Robert B., Jr. 2000. Data base for early postfire succession in Northern Rocky Mountain forests. Gen. Tech. Rep. RMRS-GTR-61-CD, [CD-ROM]. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. On file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. [43743]

210. Stuart, John D. 1987. Fire history of an old-growth forest of Sequoia sempervirens (Taxodiaceae) forest in Humboldt Redwoods State Park, California. Madrono. 34(2): 128-141. [7277]

211. Sugihara, Neil G.; Reed, Lois J. 1987. Vegetation ecology of the Bald Hills oak woodlands of Redwood National Park. Tech. Rep. 21. Orick, CA: Redwood National Park Research and Development, South Operations Center. 78 p. [55266]

212. Swedberg, Kenneth C. 1973. A transition coniferous forest in the Cascade Mountains of northern Oregon. The American Midland Naturalist. 89(1): 1-25. [4806]

213. 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.; Stewart, Ronald E.; Tappeiner, John C., II; Wells, Gail E., eds. Reforestation practices in southwestern Oregon and northern California. Corvallis, OR: Oregon State University, Forest Research Laboratory: 136-164. [22157]

214. Tappeiner, John; Zasada, John. 1988. Ecology and management of shrubs and hardwoods in Oregon's Coast Range forests. Cope Report. Newport, OR: Oregon State University, Hatfield Marine Science Center. 1(4): 3-4. [15445]

215. Tesch, Steven D.; Crawford, Michael S.; Baker-Katz, Kathryn; Mann, John W. 1990. Recovery of Douglas-fir seedlings from logging damage in southwestern Oregon: preliminary evidence. Northwest Science. 64(3): 131-139. [11764]

216. Thilenius, John F. 1968. The Quercus garryana forests of the Willamette Valley, Oregon. Ecology. 49(6): 1124-1133. [8765]

217. Thilenius, John F.; Hungerford, Kenneth E. 1967. Browse use by cattle and deer in northern Idaho. Journal of Wildlife Management. 31: 141-145. [76405]

218. Thompson, K. 1992. The functional ecology of seed banks. In: Fenner, Michael, ed. Seeds: The ecology of regeneration in plant communities. Wallingford, UK: C.A.B. International: 231-258. [60802]

219. Thompson, Ralph L. 2001. Botanical survey of Myrtle Island Research Natural Area, Oregon. Gen. Tech. Rep. PNW-GTR-507. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 27 p. [43785]

220. Thysell, David R.; Carey, Andrew B. 2000. Effects of forest management on understory and overstory vegetation: a retrospective study. Gen. Tech. Rep. PNW-GTR-488. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 41 p. [47255]

221. Topik, Christopher. 1989. Plant association and management guide for the grand fir zone, Gifford Pinchot National Forest. R6-Ecol-TP-006-88. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 110 p. [11361]

222. 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. [2351]

223. 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. [3234]

224. Turner, Nancy J.; Cocksedge, Wendy. 2001. Aboriginal use of non-timber forest products in northwestern North America: applications and issues. Journal of Sustainable Forestry. 13(3-4): 31-57. [39451]

225. U.S. Department of Agriculture, Forest Service, Division of Timber Management, Region 1. 1970. Reference material: Daubenmire habitat types. Unpublished report on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 17 p. [+ appendices]. [17399]

226. U.S. Department of Agriculture, Forest Service. 1937. Range plant handbook. Washington, DC. 532 p. [2387]

227. U.S. Department of Agriculture, Natural Resources Conservation Service. 2010. PLANTS Database, [Online]. Available: https://plants.usda.gov /. [34262]

228. U.S. Department of the Interior, Bureau of Land Management. 1993. Fire effects in sagebrush/grass and pinyon-juniper plant communities. In: Fire effects in plant communities on the public lands. EA #MT-930-93-01. [Billings, MT]: U.S. Department of the Interior, Bureau of Land Management, Montana State Office: I-1 to I-42. [55086]

229. Villarin, Lauren A.; Chapin, David M.; Jones, John E., III. 2009. Riparian forest structure and succession in second-growth stands of the central Cascade Mountains, Washington, USA. Forest Ecology and Management. 257(5): 1375-1385. [73721]

230. Vogl, Richard J. 1973. Ecology of knobcone pine in the Santa Ana Mountains, California. Ecological Monographs. 43: 125-143. [4815]

231. Waring, R. H.; Major, J. 1964. Some vegetation of the California coastal redwood region in relation to gradients of moisture, nutrients, light, and temperature. Ecological Monographs. 34: 167-215. [8924]

232. Weaver, Harold. 1968. Fire and its relationship to ponderosa pine. In: Proceedings, California Tall Timbers fire ecology conference; 1967 November 9-10; Hoberg, CA. Number 7. Tallahassee, FL: Tall Timbers Research Station: 127-149. [16903]

233. Weber, William A.; Wittmann, Ronald C. 1996. Colorado flora: eastern slope. 2nd ed. Niwot, CO: University Press of Colorado. 524 p. [27572]

234. Weisberg, Peter J. 1998. Fire history, fire regimes, and development of forest structure in the central western Oregon Cascades. Corvallis, OR: Oregon State University. 256 p. Dissertation. [40273]

235. Wells, Philip V. 1962. Vegetation in relation to geological substratum and fire in the San Luis Obispo quadrangle, California. Ecological Monographs. 32(1): 79-103. [14183]

236. Welsh, Stanley L.; Atwood, N. Duane; Goodrich, Sherel; Higgins, Larry C., eds. 1987. A Utah flora. The Great Basin Naturalist Memoir No. 9. Provo, UT: Brigham Young University. 894 p. [2944]

237. Wender, Bryan W.; Harrington, Constance A.; Tappeiner, John C., II. 2004. Flower and fruit production of understory shrubs in western Washington and Oregon. Northwest Science. 78(2): 124-140. [51134]

238. White, Diane E.; Atzet, Thomas; Martinez, Patricia A. 2004. Relationship of historic fire regimes to dead wood components in white fir forests of southwestern Oregon. In: Engstrom, R. Todd; Galley, Krista E. M.; de Groot, William J., eds. Fire in temperate, boreal, and montane ecosystems: Proceedings of the 22nd Tall Timbers fire ecology conference: an international symposium; 2001 October 15-18; Kananaskis Village, AB. No. 22. Tallahassee, FL: Tall Timbers Research, Inc: 117-124. [52307]

239. Whitlock, Cathy; Skinner, Carl N.; Bartlein, Patrick J.; Minckley, Thomas; Mohr, Jerry A. 2004. Comparison of charcoal and tree-ring records of recent fires in the eastern Klamath Mountains, California, U.S.A. Canadian Journal of Forest Research. 34(10): 2110-2121. [55446]

240. Whittaker, R. H. 1953. A consideration of climax theory: the climax as a population and pattern. Ecological Monographs. 23(1): 41-78. [64492]

241. Wiens, John A.; Nussbaum, Ronald A. 1975. Model estimation of energy flow in northwestern coniferous forest bird communities. Ecology. 56: 547-561. [19167]

242. Wittinger, W. T.; Pengelly, W. L.; Irwin, L. L.; Peek, J. M. 1977. A 20-year record of shrub succession in logged areas in the cedar-hemlock zone of northern Idaho. Northwest Science. 51(3): 161-171. [6828]

243. Wolter, B. H. K.; Fonda, R. W. 2002. Gradient analysis of vegetation on the north wall of the Columbia River Gorge, Washington. Northwest Science. 76(1): 61-76. [65993]

244. Wright, Henry A. 1978. The effect of fire on vegetation in ponderosa pine forests: A state-of-the-art review. Lubbock, TX: Texas Tech University, Department of Range and Wildlife Management. 21 p. In cooperation with: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. [4425]

245. Wright, Henry A.; Neuenschwander, Leon F.; Britton, Carlton M. 1979. The role and use of fire in sagebrush-grass and pinyon-juniper plant communities: A state-of-the-art review. Gen. Tech. Rep. INT-58. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 48 p. [2625]

246. Yerkes, Vern P. 1960. Occurrence of shrubs and herbaceous vegetation after clear cutting old-growth Douglas-fir. Res. Pap. PNW-34. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 12 p. [8937]

247. Young, J. A.; Hedrick, D. W.; Keniston, R. F. 1967. Forest cover and logging--herbage and browse production in the mixed coniferous forest of northeastern Oregon. Journal of Forestry. 65: 807-813. [16290]

248. Young, Richard P. 1983. Fire as a vegetation management tool in rangelands of the Intermountain region. In: Monsen, Stephen B.; Shaw, Nancy, comps. Managing Intermountain rangelands--improvement of range and wildlife habitats: Proceedings of symposia; 1981 September 15-17; Twin Falls, ID; 1982 June 22-24; Elko, NV. Gen. Tech. Rep. INT-157. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: 18-31. [2681]

249. Youngblood, Andrew; Metlen, Kerry L.; Coe, Kent. 2006. Changes in stand structure and composition after restoration treatments in low elevation dry forests of northeastern Oregon. Forest Ecology and Management. 234(1-3): 143-163. [64992]

250. Youngblood, Andrew; Wickman, Boyd E. 2002. The role of disturbance in creating dead wood: insect defoliation and tree mortality in northeastern Oregon. In: Laudenslayer, William F., Jr.; Shea, Patrick J.; Valentine, Bradley E.; Weatherspoon, C. Phillip; Lisle, Thomas E., tech. coords. Proceedings of the symposium on the ecology and management of dead wood in western forests; 1999 November 2-4; Reno, NV. Gen. Tech. Rep. PSW-GTR-181. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 155-168. [44351]

251. Zack, Arthur C.; Morgan, Penelope. 1994. Early succession on two hemlock habitat types in northern Idaho. In: Baumgartner, David M.; Lotan, James E.; Tonn, Jonalea R., compilers. Interior cedar-hemlock-white pine forests: ecology and management: Symposium proceedings; 1993 March 2-4; Spokane, WA. Pullman, WA: Washington State University, Department of Natural Resources: 71-84. [25792]

252. Zimmerman, G. T.; Neuenschwander, L. F. 1984. Livestock grazing influences on community structure, fire intensity, and fire frequency within the Douglas-fir/ninebark habitat type. Journal of Range Management. 37(2): 104-110. [10103]

253. Zimmerman, Gordon Thomas. 1979. Livestock grazing, fire, and their interactions within the Douglas-fir/ninebark habitat type of northern Idaho. Moscow, ID: University of Idaho. 145 p. Thesis. [6724]

254. Zinke, Paul J. 1977. The redwood forest and associated north coast forests. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley and Sons: 679-698. [7212]

255. Zobel, Donald B. 2002. Ecosystem use by indigenous people in an Oregon coastal landscape. Northwest Science. 76(4): 304-314. [64344]

256. 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. [8767]

257. Zobel, Donald B.; Roth, Lewis F.; Hawk, Glenn M. 1985. Ecology, pathology, and management of Port-Orford-cedar (Chamaecyparis lawsoniana). Gen. Tech. Rep. PNW-184. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 161 p. [9245]