Sambucus racemosa



INTRODUCTORY


 

Red elderberry along Turnagain Arm near Bird, Alaska. Photo by Mary Hopson.

AUTHORSHIP AND CITATION:
Fryer, Janet L. 2008. Sambucus racemosa. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].

FEIS ABBREVIATION:
SAMRAC
SAMRACR
SAMRACM

NRCS PLANT CODE [233]:
SARA2
SARAM4
SARAR3

COMMON NAMES:
red elderberry
red elder
stinking elderberry

TAXONOMY:
The scientific name of red elderberry is Sambucus racemosa L. (Caprifoliaceae) [57,95,118,126,135,151]. There are 2 North American varieties:

Sambucus racemosa L. var. racemosa, typical variety of red elderberry [135]
Sambucus racemosa L. var. melanocarpa (A. Gray) McMinn [57,81,121,135,151,240], Rocky Mountain elderberry

In this review, "red elderberry" refers to the species. Varieties are referred to as either the "typical variety" or "Rocky Mountain elderberry".

Natural hybridization between red elderberry and other Sambucus species is apparently rare [132]. Degree of hybridization between red elderberry varieties was not described in the literature as of 2008.

SYNONYMS:
forSambucus racemosa L. var. melanocarpa:
  Sambucus melanocarpa A. Gray [137,186]

forSambucus racemosa L. var. racemosa:
  Sambucus callicarpa Greene [236]
  Sambucus microbotrys Rydb. [121,137,186,239]
  Sambucus pubens Michx. [81,194,218]
  Sambucus pubens Michx. var. arborescens Torr. & A. Gray [204,244]
  Sambucus racemosa L. var. arborescens (Torr. & A. Gray) A. Gray [126]
  Sambucus racemosa L. var. leucocarpa (Torr. & A. Gray) Cronquist
  Sambucus racemosa L. var. laciniata W.D.J. Koch ex DC.
  Sambucus racemosa L. var. microbotrys (Rydb.) Kearney & Peebles [57,118,137,240]
  Sambucus racemosa L. var. pubens (Michx.) Koehne [95,121]
  Sambucus racemosa L. subsp. pubens (Michx.) House [98,126,178,209]

LIFE FORM:
Shrub-tree

FEDERAL LEGAL STATUS:
None

OTHER STATUS:
Information on state- and province-level protection status of red elderberry and its varieties in the United States and Canada is available at NatureServe.

DISTRIBUTION AND OCCURRENCE

SPECIES: Sambucus racemosa
GENERAL DISTRIBUTION:
Red elderberry is native to North America and Eurasia [57,95,126,137]. In North America it occurs from Alaska and Yukon south to California, New Mexico, South Dakota, Missouri, and Georgia and east to Quebec, Newfoundland, Maine, and North Carolina. The typical variety occurs throughout the range of the species. Rocky Mountain elderberry occurs from British Columbia and Alberta south to California, New Mexico, and North Dakota, overlapping the distribution of the typical variety in those areas [135]. Plants Database provides a distributional map of red elderberry and its varieties.

HABITAT TYPES AND PLANT COMMUNITIES:
Red elderberry is common in many forest communities but is rarely dominant (for example, [30,61,62,84,183,202,232]). It typically occurs in scattered patches or as isolated individuals [232], although it often forms thickets in northern Utah [52]. Red elderberry grows in riparian zones across its distribution (for example, [22,162,174,190,201,204]).

Red elderberry is common in conifer series in the West (contiguous United States west of the Mississippi River), particularly fir-spruce (Abies-Picea spp.) and other mesic or wet forest types [65,69,82,161]. It is a frequent associate in red alder (Alnus rubra) communities of the Pacific coast [114].

Red elderberry is also a common component of red alder and other shrubfields in Alaska [112]. It grows in Sitka alder-willow (Alnus viridis subsp. sinuata-Salix spp.) communities in south-central Alaska [176].

In the Great Lakes and eastern regions, red elderberry occurs in hardwood [34,38,61], mixed, and spruce-fir ([30], review by [63]) forests. It may be one of only a few woody species to persist in the understory of closed-canopy, old-growth eastern forests [38]. In a floristics study of northern mixed-hardwood communities, relative abundance of red elderberry increased from southern Michigan to east-central Minnesota [202,203].

Vegetation classifications describing plant communities in which red elderberry is a dominant or indicator species are listed below.

Idaho Montana Utah Wisconsin British Columbia Quebec

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Sambucus racemosa

 

National Park Service photo by Richard Thomas.

GENERAL BOTANICAL CHARACTERISTICS:
Botanical description: This description provides characteristics that may be relevant to fire ecology and is not meant for identification. Keys for identification are available (for example, [57,118,121,218,238,239,240]).

Aboveground description: Red elderberries are typically low-growing shrubs [186,236] but may grow 7 to 20 feet (2-6 m) m) tall [118,209]. They may grow as single shrubs or trees [121,179,236] or form clumps [179,236] or thickets [52]. Red elderberries have pubescent, pithy, soft to barely woody branches [102,209,218] with soft bark [126]. Red elderberry is deciduous [98,236]. The large, compound leaves are opposite, with ovate-lanceolate leaflets that are downy on their undersides. The inflorescence is a large, showy, panicled cyme [218,218,236,240] bearing small (~3 mm long), numerous flowers [126,209,218]. The fruits are berrylike drupes [209,236,240]. At maturity, drupes of the typical variety are red [57,240], or rarely, yellow or white [137,238], while those of Rocky Mountain elderberry are purplish-black to black [57,102,240]. The seeds are nutlets [57,98], with 2 to 4 seeds/fruit [228]. The foliage, branches, and flowers are foul-scented when crushed [126,209].

Belowground description: Underground structure of red elderberry was described in few studies as of 2008. An Ontario study found red elderberry had highly branched lateral roots, with diameter of secondary roots ranging from 1.38 to 2.76 inches (3.50-7.00 cm). Maximum length of root hairs (1.6 inches (4.0 cm)) was low relative to associated species [33]. The typical variety of red elderberry is sometimes rhizomatous ([52,241],  review by [102]); however, on the West coast, the typical variety may lack rhizomes (Newton 1984, personal communication cited in [102]). As of 2008, it was unclear how often rhizomes occurred within and/or among red elderberry populations. A northern Utah study found red elderberry rhizomes and roots extended 3 feet (1 m) deep [52]. Area of red elderberry clones in the Rocky Mountains may range from 1 to 16 m² [102].

Raunkiaer [197] life form:
Phanerophyte
Geophyte (applies only to rhizomatous populations)

REGENERATION PROCESSES:
Red elderberry reproduces vegetatively and from seed (review by [245]).

Vegetative regeneration: Red elderberry sprouts from the root crown and/or rhizomes after top-kill ([28,52],  reviews by [25,26,102,245]). It also layers [94,245]. Sprouting is generally the most common method of red elderberry regeneration, although seedling establishment allows for colonization in new areas [245]. Red elderberry seedlings were rarely observed, for example, over 3- to 5-year periods in quaking aspen forest-shrubland mosaics of northern Utah, although red elderberry clones were abundant in the area [52].

Pollination and breeding system: Red elderberry is outcrossing [138]. Flowers are pollinated by bees, flies [138], and wind [132].

Flower, fruit, and seed production: Red elderberry usually produces a good seed crop every year [102,179]. On the Monongahela National Forest, West Virginia, 70% to 80% of red elderberry plants bore fruit in 4 consecutive years [191]. In northern Utah, 40% of aerial red elderberry stems produced flowers. The number of flowers/inflorescence ranged from 100 to 400, with 60% to 90% of flowers producing fruit. Approximately 30% of seeds were unfilled; filled seeds were 74% to 91% viable in the laboratory [52]. On a study on the Olympic Peninsula of Washington, only 2 of 52 red elderberry shrubs failed to produce flowers and fruit. Production by the remaining 50 shrubs was [241]:

Mean (and range) of reproductive and growth characteristics for red elderberries in Washington [241]
Number of flowers/stem Shrub age  Basal diameter (cm) Height (m) Number of stems/shrub
1,0875 (144-17,064) 6 (2-13) 3.4 (1.1-9.4) 3.2 (1.5-5.8) 3 (1-7)

Flower and fruit abundance was positively associated with plant DBH and canopy gaps and negatively associated with overstory density (P=0.05). Red elderberries with greatest fruit production were found in canopy gaps more frequently than under canopies [241].

Closed canopies or browsing can reduce red elderberry flower and fruit production. A study of fleshy-fruited wildlife shrubs on the Kenai Peninsula of Alaska showed canopy cover significantly decreased red elderberry fruit production (P=0.01) [220]. On the Ottawa National Forest, Michigan, white-tailed deer browsing significantly reduced the number of flowers and fruits on red elderberries (P<0.05) [148]. In a Sitka spruce-western hemlock (Tsuga heterophylla) forest in Olympic National Park, Washington, cover of red elderberry and other shrub species was significantly greater inside than outside elk and mule deer exclosures (P<0.05). Refugia patches that were inaccessible to ungulates, such as exposed root mats of large, fallen conifers and areas behind stacks of fallen logs, allowed red elderberry to bear flowers and/or fruit. Red elderberries in other areas were browsed too heavily to have "appreciable" flower and fruit production [207].

Seed dispersal: Frugivorous birds and mammals disperse red elderberry seeds [35,81,102,120,150,219]. Plant location influences the likelihood of animal frugivory and seed dispersal. In British Columbia, passerine birds deposited significantly more red elderberry seeds in the understory of a Sitka spruce-hemlock forest than at forest edges (P<0.01) [35]. In Wisconsin, frugivorous birds selected significantly more red elderberry fruits from isolated plants than from clustered plants. Large fruit size, heavy fruit production, and fruit sweetness increased the probability of fruit selection and seed dispersal for individual plants (P<0.01 for all measures) [70].

Water- and/or gravity-dispersed seed may be unimportant to red elderberry establishment. In a study using seed from a mixed-hardwood community by Little Otter Creek in Vermont, 100% of red elderberry germinants emerging in the greenhouse came from soil samples; no red elderberry germinants emerged from floodplain debris samples or seed rain traps [124].

Seed banking: Red elderberry has a litter [104] and soil [4,25,104,107,140,145] seed bank. Fallen logs may also be a seedbed for red elderberry [173]. Red elderberry is often strongly represented in the soil seed bank. Behind red alder, seeds of red elderberry were the second most common in soils beneath a red alder community in British Columbia. Viable red elderberry seeds were distributed in the top 4 inches (10 cm) of soil [139]. Red elderberry seeds were "especially common" (frequency of at least 50%) in the soil seed banks of Douglas-fir (Pseudotsuga menziesii) and grand fir forests on the Payette and Boise National Forests of Idaho. Density varied from 0 to 210 red elderberry seeds/m². Seeds were found mostly in upper soil layers (0.4 inch (5 cm) deep), although a few seeds were found at depths to 17 inches (43 cm). In the greenhouse, mean germination of the soil-stored seed was 16% [149]. A study of the seed bank beneath an eastern hemlock forest in New York found red elderberry was 1 of only 4 woody species with viable seeds in the soil; all other seeds were from herbaceous species [247].

In a greenhouse experiment using soil collected from different-aged, mixed-hardwood stands on the White Mountain National Forest, New Hampshire, red elderberry showed variable density in the soil seed bank [97]. Density of viable seed was apparently not tied to stand age:

Density of red elderberry germinants in forest floor samples from mixed-hardwood stands in New Hampshire [97]

Stand age 

5 38 95 200+
Seeds/m² 42 25 3 69

Red elderberry's representation in the soil seed bank varied in 2 studies comparing logged and unlogged sites. In coastal British Columbia, red elderberry seed was present in soil samples from an unlogged, old-growth western redcedar (Thuja plicata)-western hemlock forest but not in samples from an adjacent logged forest [140]. However, a field study using potted plants on the Hubbard Brook Experimental Forest, New Hampshire, had opposite results. No red elderberry germinants emerged from soil collected from an unlogged sugar maple-American beech-yellow birch (Betula alleghaniensis) forest, but red elderberry density was 9.2 germinants/m² in soil collected from an adjacent logged forest [125].

Red elderberry is represented only in the seed bank on some sites. In a balsam fir (A. balsamifera) forest type in the Boundary Waters Canoe Area of Michigan, red elderberry was present in the seed bank at a mean density of 484,000 seeds/ha but was not present in aboveground vegetation. Spruce budworms had killed most of the overstory trees [4]. A study on the Olympic Peninsula of Washington had similar findings: red elderberry plants were not present in a closed-canopy Douglas-fir-western hemlock forest, but in the greenhouse, red elderberry germinants emerged from litter and soil samples collected in the forest [104].

Germination: Red elderberry seeds are dormant ([29], review by [20], Pelton 1956, cited in [17]). Embryological dormancy [20,29,120] and a hard seedcoat maintain seed dormancy [20,29]. Fire, the digestive acids of frugivorous animals, or stratification can break dormancy. Seedcoat damage from fire or digestion may speed up and/or increase germination rates. Heat from fire can break dormancy by cracking the hard seedcoat [25,26,208]. An Alaskan experiment found that red elderberry seeds showed faster and higher germination rates after passage through the digestive tracks of passerines or American black bears (P=0.0001). Animal species influenced germination: 31% of red elderberry seeds ingested by varied thrushes germinated, while only 19% of seeds ingested by American robins and 14% of seeds ingested by American black bears germinated [228]. In the laboratory, damaging the seedcoat by clipping or acid bath, warm stratification, cold stratification, and warm stratification followed by cold stratification have broken dormancy [20,29,120,245]. A laboratory experiment by Hidayati and others [119,120] found that red elderberry seeds did not require treatments that damaged the seedcoats to imbibe water. A greenhouse study found light improved germination rates of seeds given warm stratification followed by cold stratification (P=0.05). In an unheated greenhouse, germination rate of fresh red elderberry seed was nearly 100% after overwintering [120]. A wide range of germination (6-100%) is reported for red elderberry in laboratory experiments [20,102]. Red elderberry may require 2 years to complete germination in the field [29]. Open, disturbed soil provides a favorable seedbed. Red elderberry establishment is also reported on decayed wood [102].

See Immediate fire effect on plant for further information on the possible effects of fire to red elderberry seed.

Seedling establishment and plant growth: Red elderberry seedlings establish best on open, disturbed sites [52]. They do not establish well beneath parent plants (Newton 1984, personal communication cited in [102]) or closed forests. In an Upper Michigan study comparing seed bank and seedling populations in forest gaps and closed-canopy sugar maple-eastern hemlock forest, red elderberry seedlings were present in low numbers in forest gaps but did not occur in closed stands. In the greenhouse, significantly more red elderberry germinants emerged from soil collected in gaps compared to soil beneath mature forest [177]. See Gap succession for more information on the importance of gaps to red elderberry.

Mean frequency and density of red elderberry seedlings and germinants emerging from the seed bank in Upper Michigan [177]
Age class Frequency (%) Density (stems/0.01 ha)
Gap Forest Gap Forest
Seedlings 2.2 not present 3.30 not present
Germinants from seed bank 44.4 30.0 93.9* 45.3
*Significant difference between gap and forest plots (P<0.001).

Red elderberry sprouts typically grow faster than seedlings, with sprout growth rates of 10 to 15 feet (3-4.5 m) in their first year (Newton 1984, personal communication cited in [102]). Rhizome sprouts may flower in their 2nd year [52]. Red elderberry seedlings may grow rapidly on favorable sites, however. Seedlings have produced a full crown and reached 10 feet (3 m) tall in 3 to 4 years on sites in the Intermountain West [102]. On the Oregon coast, 4-year-old red elderberry seedlings were 1 to 3 feet (0.3-0.9 m) tall [49]. Seedlings in the greenhouse have produced rhizomes by age 3 [102].

SITE CHARACTERISTICS:
Red elderberry occurs in thickets, woodlands, forests, and subalpine meadows [57,65,118,126,209,218]. Site characteristics vary widely across red elderberry's broad geographic and elevational range.

Climate, moisture, and nutrient regime: Red elderberry grows in interior and coastal climates but is rare in arid regions. On the Kenai Peninsula, red elderberry was positively correlated with the mesic climate of interior lowlands but not the moist, maritime climate of the coast (P<0.05) [24]. In the Rocky Mountains, Davis [65] reported a mean of 34 inches (860 mm) annual precipitation for subalpine red elderberry shrubfields and spruce-fir forests with a red elderberry component. Red elderberry is apparently less tolerant of warm climates than blue elderberry (Sambucus nigra subsp. cerulea), and is confined to relatively cool, moist riparian zones, swamps, and mountainous areas in its southern distribution [245]. In Arizona, red elderberry is reported only from the San Francisco Peaks and the Grand Canyon [137].

Red elderberry prefers moist, [98,102,236], nutrient-rich [143,150] sites but occasionally grows in dry areas. It is often found in riparian zones [22,179,204], wet meadows, and moist to wet parklands [179], and is considered a facultative wetland species [147]. In montane zones, it is most common in openings where snow accumulates and remains until late spring [179,195]. Red elderberry is reported on dry to very moist, nutrient-rich soils in British Columbia [143,150].

Red elderberry can be an indicator species for site productivity. A study on the Central Coast Range of Oregon found red elderberry was positively correlated with site productivity (P<0.05) [41], and red elderberry is an indicator of moderately to richly productive sites in Wisconsin [146]. A study on the Kenai Peninsula of southeastern Alaska also found red elderberry was positively correlated with site productivity (P<0.05) [24].

Red elderberry is moderately flood tolerant. In the Fraser River valley of British Columbia, red elderberry died back to the root crown and sprouted after a 50-year flood in 1948 [28].

Soil properties and parent materials: Red elderberry generally grows in deep, well-drained, loamy soils [52,102,245]. It is reported on sandy loam and loam in the Great Lakes states [48]. Soil pH across red elderberry's range varies from acidic [242] to basic, although neutral soils are preferred [245]. Red elderberry grows on limestone-derived soils along Lake Champlain; anorthosite and gneiss parent materials in the Adirondack Mountains; and on shale, sandstone, and conglomerate materials in the Catskill Mountains [150].

Elevation: Red elderberry generally grows in submontane to montane zones in the West [151], although its range extends into alpine fellfields in California [186]. Red elderberry occurs mostly in high-elevation montane zones in the East [194,244].

Elevational ranges across red elderberry's distribution. Information pertains to the species unless a variety is specified.
Arizona typical variety from 7,500-10,000 feet; Rocky Mountain red elderberry found ≥7,500 feet [137]
California <11,000 feet [118]
Colorado ≥10,000 feet in Custer County [47]
Nevada 6,200-9,000 feet [136]
Utah 4,460-1,025 feet [240]
Virginia, Blue Ridge Mountains >3,000 feet in yellow birch boulderfields [85]
Adirondack Mountains 100-3,830 feet [150]
Intermountain West 5,900-10,000 feet [57]
Rocky Mountains <3,300 feet [81]

SUCCESSIONAL STATUS:
Red elderberry prefers open sites [102,150,190] but tolerates shade [144,150]. It responds well to release [41,102].

Primary succession: Red elderberry is a pioneer species on the islands of Barkley Sound, British Columbia, where northwestern crows disperse red elderberry seeds along the rocky shorelines. In this environment, red elderberry shrubs tended to associate mostly with other red elderberries. Of 6 fleshy-fruited shrubs, red elderberry was the only species where abundance was negatively associated with presence of other shrub species (P=0.001) [37]. Red elderberry joins seral thicket formations in rockshore succession on the islands of Lake Michigan, probably establishing from bird-dispersed seed. Due to edaphic conditions on the Gull Islands and other small, rocky islands of Lake Michigan, most shrub thickets do not succeed to climax jack pine-black spruce (Pinus banksiana-Picea mariana) forest, although this occurs on larger islands such as Isle Royale [53].

Cooper [54,55] found red elderberry in the understories of willow-Sitka alder (Salix spp.-Alnus viridis subsp. sinuata) communities [54] and young Sitka spruce stands [55] in fjords of Glacier Bay, Alaska. In postglacial succession there, plant community development generally proceeds from mixed shrub/herb/moss to willow-Sitka alder, young Sitka spruce forest, and mature Sitka spruce-western hemlock forest communities [55].

Secondary succession: Red elderberry is most common in early seral communities but may occur in all stages of succession. For example, red elderberry is typically present in "large numbers" in seral pin cherry (Prunus pensylvanica) stands, which are usually maintained by frequent fire [96]. Several experts list red elderberry as a pioneer in second-growth Sitka spruce-western hemlock forests of southeastern Alaska (review by [225]). In a mixed conifer-hardwood community at the Harvard Forest of Massachusetts, red elderberry frequency on permanent plots was greatest (15%) 10 years after a hurricane. It was not found on plots 1 year before, 2 years after, or 53 years after the hurricane [166].

Seral occurrence: Rocky Mountain elderberry is a common understory component of red alder shrublands, which are usually seral to Douglas-fir and/or Sitka spruce [89,114]. Franklin and Dyrness [88] report a "strong tendency" for development of dense, pure red elderberry or mixed red alder-red elderberry shrubfields after fire or logging in the Sitka spruce zone of Oregon and Washington. Red elderberry replaces red alder successionally on some sites. On the Central Coast Range of Oregon, red elderberry established and persisted in red alder stands, releasing when the red alder became senescent [41]. In an inventory of plant community composition in different stages of floodplain succession on Vancouver Island, red elderberry was present in the young seral stage, which was dominated by red alder. It was absent from new gravel bars and mature Sitka spruce forest [46]. However, in a chronosequence study of a red alder/salmonberry community in western Oregon, red elderberry cover and frequency were variable across 51 years of succession. Red elderberry was absent from stands less than 4 years old; highest red elderberry coverage (11%) and frequency (26%) was in 49-year-old stands [117]. In 1979, Thilenius [226] found red elderberry in seral red alder communities on the Copper River Delta. The area had been uplifted by the 1964 earthquake and was changing successionally from herbland to shrubland [226].

On the Wasatch Plateau of central Utah, red elderberry was one of several woody species pioneering on depleted mountain meadow rangelands that had been subject to severe flooding after decades of overgrazing by livestock [79].

Logging: Tree harvest may favor red elderberry growth by opening the canopy, although red elderberry abundance may not increase substantially after logging (for example, [196]). Red elderberry may sprout and/or colonize from seed after tree harvest [103]. In a sugar maple-American beech-yellow birch (Betula alleghaniensis) forest on the Hubbard Brook Experimental Forest, New Hampshire, red elderberry was more frequent on logged sites for at least 3 years after logging (4.9% frequency) than on adjacent unlogged forest (<1% frequency) [125]. In a subalpine fir/queencup beadlily (Abies lasiocarpa/Clintonia uniflora) habitat type in northwestern Montana, red elderberry cover was higher on logged plots (x=60%) than on old-growth plots (x=23.5%) [248]. It was an early-seral species in cutover grand fir habitat types of central Idaho [92] and in second-growth, logged hardwood forests of Michigan [78]. Clearcutting or clearcutting followed by slash burning apparently had no effect on red elderberry abundance in western hemlock/Oregon boxwood forests of northern Idaho. Red elderberry was absent or had less than 3% cover on unlogged sites, and red elderberry cover did not increase after logging or logging and burning [243].

Logging method can affect the rate of red elderberry recovery after tree harvest, although trends in red elderberry response to different logging methods are not well studied. On the Fort Lewis Military Reservation, Washington, red elderberry was not present on Douglas-fir sites that had been harvested in the 1930s and then left unmanaged for 70 years, but red elderberry had 22.5% frequency and 2.2% cover on nearby, thrice-thinned Douglas-fir sites, showing a significant difference between sites (P<0.001). Red elderberry was considered an indicator of cutover or disturbed sites [227]. Heavy logging may favor red elderberry on some sites. Red elderberry attained a density of 25 stems/ha 20 years after clearcutting in a balsam fir-yellow birch forest in eastern Quebec [11] and was among the 10 most common shrubs after clearcutting in a red fir (A. magnifica var. magnifica) forest in California [16]. In a sugar maple-yellow birch forest in Upper Michigan, the mean importance value of red elderberry 2 years after logging was greater in clearcuts than in group selection or unlogged sites. Fifty years after treatment, however, red elderberry was more important in group selection cuts than on clearcuts. The 50-year importance value on unlogged sites was not calculated due to a blowdown on that site 26 years after treatments [175].

Logging sometimes reduces red elderberry abundance. Thinning to remove small-diameter trees in a Sitka spruce-western hemlock forest in west-central Oregon resulted in loss of red elderberry plants. Although present before thinning, red elderberry was absent from study plots 17 years later [5]. It was negatively associated with logged black spruce forests of northeastern Ontario [32].

Late succession: Red elderberry may be present in the understory of mature or climax forests. It was a component of the shrub layer in a climax mixed-hardwood forest in the Black Mountains of North Carolina [66] and was listed as one of the most common species in mature white spruce (Picea glauca) forests of interior British Columbia [77]. Since red elderberry is only moderately shade tolerant [144,150], gap succession helps maintain red elderberry in late stages of forest succession.

Gap succession: Red elderberry commonly establishes in canopy gaps within mature forests. It grew in openings after woolly adelgid attacks on mature balsam firs (Abies balsamifera) in Great Smoky Mountains National Park and the Pisgah National Forest, North Carolina. Red elderberry attained sapling size within 7 to 15 years after the attacks [72]. On the Hemlock Hill Biological Research Station in Pennsylvania, red elderberry established from bird-dispersed seeds in gaps within a mixed-hardwood forest. Scarlet tanagers were the primary seed dispersers [219]. In southeastern Alaska, red elderberry seedlings established in windthrow gaps in an old-growth western hemlock forest [6]. In northern Utah, the typical variety of red elderberry was most abundant in canopy openings but also grew beneath quaking aspen canopies [52].

SEASONAL DEVELOPMENT:
Animals disperse red elderberry seeds in late summer or fall. The seeds typically germinate in spring after overwintering [120], although seeds dispersed or sown in late fall may not emerge until their second spring [245]. In an unheated greenhouse, red elderberry seeds sown in August mostly emerged the next spring—from 21 to 28 March—although a few seeds germinated that fall, in late October and early November [120]. In cold climates, red elderberry plants die back to the root crown in winter [72,179]. Red elderberry begins vegetative growth early in the growing season. In the Rocky Mountains, red elderberry initiates stem bud growth while snow is still on the ground, with stem elongation completed 3 to 4 weeks after snowmelt [52]. In a sugar maple forest in Ontario, red elderberry leafed out in early spring before associated tree species [33]. Red elderberries also begin flowering relatively early in spring, when their leaves are unfolding [81]. Red elderberry is an early-fruiting tree relative to most associated woody species [36,71]. Fruits ripen from early (Bailey 1906, cited in [120]) to late [236] summer, depending on location. Fruits within panicles, on the same tree, and within populations tend to ripen synchronously [70,71].

Red elderberry phenology by state or region. Information pertains to the species unless a variety is specified.

Area Event
Alaska flowers May-July; fruits July-August [236]
Arizona typical variety flowers June-July; Rocky Mountain red elderberry flowers May-July [137]
California June-August [186].
Carolinas flowers late April-early June; fruits late June-August [194]
Colorado Rocky Mountain elderberry flowers through late spring [141]
Idaho, northern May-July [192]
Illinois flowers June-July [178]
Michigan flowers in spring; fruits in early to late summer [238]
Nevada flowers May-June [136]
Ohio flowers in May; fruits June-November [44]
western Oregon fruits late June-mid-August [130]
Pennsylvania fruits late June-early July [219]
Utah, Wasatch Mountains seeds germinate March-early April; rhizome buds of mature plants expand late May-early June beneath snow; rhizome sprouts emerge at snowmelt (~10 June); flowers emerge mid-June-early July; fruits green ~7 July; fruits and seeds mature late July-mid-August [52]
Wisconsin fruits ripens mid-July [70,71]
Northeast May-June [95]
Washington flowers April-July; seeds disperse June-July [241]
West Virginia flowers April-May; fruits June-August [218]
     Monongahela National Forest fruits 12 September-5 October [191]
Adirondack Mountains flowers in bud 4-5 May; flowers 17 May-2 June; fruits 13 July-8 August [150]
Blue Ridge Mountains April-June [244]
Intermountain West flowers June-July [57]
Great Plains flowers May-June [98]
Northeast flowers April-May [245]; fruits June-November [81,217,245]
Pacific Northwest typical variety: flowers March-July [121]; fruits June-September [102,199]; ripening may extend to December with mild winters [199]
Rocky Mountain elderberry: flowers May-June; fruits June-September [102,121,199], but ripening may extend to December with mild winters [199]
British Columbia flowers in May; fully leafed out in mid-June; fruits in June [102]
     Vancouver Island fruits 9 July-18 August [36]
Nova Scotia flowers 1-20 June [204]
Ontario flowers May-June; fruits in midsummer [209]
     Great Lakes-St Lawrence region leafs out 6-28 April; roots expand early spring-early summer; flowers in late spring; leaves senesce in late fall; roots senesce from late summer-early fall [33]

FIRE EFFECTS AND MANAGEMENT

SPECIES: Sambucus racemosa
FIRE EFFECTS: Immediate fire effect on plant: Fire can crack red elderberry's hard seedcoat [25,26,208], which may enhance germination [25,26]. Fire typically top-kills red elderberry plants [25,26,165,171,215]. Stickney [215] rates red elderberry as low in susceptibility to fire kill because perennating buds on red elderberry's root crown are protected by mineral soil.

Prolonged heat kills red elderberry seed. A greenhouse study found red elderberry showed better seedling emergence when not subject to sustained high temperatures. Soil samples were collected from an unburned subalpine fir/big huckleberry (Vaccinium membranaceum) forest in Yellowstone National Park, Wyoming. After 6 months in the greenhouse, unheated soil samples had a mean density of 40 red elderberry germinants/m², while soil samples heated to 120 °F (50 °C) for 1 hour had a mean density of 13 germinants/m². Red elderberry did not emerge from soil samples heated to 212 °F (100 °C) or 302 °F (150 °C) for 1 hour [45].

Postfire regeneration strategy [165,214,216]:
Tree with a sprouting root crown
Tall shrub with a sprouting root crown
Rhizomatous shrub, rhizome in soil (applies to only rhizomatous populations)
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: Red elderberry sprouts from the root crown and/or rhizomes after top-kill by fire ([25,26,165,171,215], Newton 1984, personal communication cited in [102]). Not all red elderberry populations are rhizomatous (see Belowground description), so postfire rhizome sprouting will not occur on all sites.

Red elderberry may establish after fire from on-site seed stored in litter [107] or soil [100,107,108,109,216] or from off-site seed. Red elderberry seeds in the seed bank may begin germinating the spring after fire (see Regeneration Processes and Seasonal Development) [25,26,208]. Because red elderberry seed is animal dispersed, postfire establishment from off-site seed sources is likely; however, animal dispersal onto burns was not well documented in the literature as of 2008. A year after the severe Sundance Wildfire in northern Idaho, red elderberry seedlings and root crown sprouts on study sites had 3% cover and 4% frequency (seedling and root crown sprout data were pooled) [213].

Red elderberry does not often gain dominance after fire; it typically remains a minor component of the vegetation on sites where it occurred in low numbers before fire (for example, [164]). However, postfire seedling emergence may be "extensive" on some sites [25,26,208].

Many studies show red elderberry is favored but not greatly enhanced by fire [102,106,110,152,181,183,212,246]. In western Montana, red elderberry attained minor coverage (0.2-0.3%) 2 years after both the Miller Creek prescribed fire on the Flathead National Forest and the Newman Ridge prescribed fire on the Lolo National Forest. Red elderberry was not present on study sites before the fires [212], suggesting that it established from on- or off-site seed, not sprouts. After spring wildfires in a sugar maple-eastern hemlock-American beech forest in south-central New York, red elderberry saplings reestablished from sprouts but had the lowest importance value (2.5) of all woody species present [221]. Red elderberry had 0.3% cover 35 years after a wildfire in a red spruce-Fraser fir (Picea rubens-Abies fraseri) forest in North Carolina. Its density at postfire year 35 was estimated at 1,364 stems/ha [206]. In chronosequence studies in New Brunswick, red elderberry was present in jack pine and mixed-hardwood forests that developed 7 to 20 years after stand-replacing fires. Red elderberry was present in mostly trace amounts. Its cover (3%) and frequency (3%) were greatest in 10-year-old burns [167].

On many sites, a combination of fire and other treatments may have little effect on red elderberry abundance.

Logging and burning: After clearcutting and slash burning in subalpine fir-western larch-Engelmann spruce (Larix occidentalis-Picea engelmannii) stands on the Flathead National Forest, Montana, red elderberry had a nonsignificant, 5.6% mean decrease in frequency compared to unburned stands. Time-since-fire on study sites ranged from 2 to 15 years [237].

Red elderberry responses to winter clearcutting and slash burning were similar in studies on the Clearwater, MacKenzie, and Headwaters Forest Districts of south-central, central interior, and east-central British Columbia, respectively. All sites were logged when snow was deep enough to prevent disturbance to understory vegetation or the forest floor [108,109,111]. On the Clearwater Forest District, red elderberry established from on-site seed after winter clearcutting of an Engelmann spruce-subalpine fir forest followed by spring or fall slash burning, with red elderberry frequency consistently higher in burned than in unburned plots (P<0.01). Red elderberry cover was generally low. In postfire year 2, a flush of postfire red elderberry germination resulted in greater cover on burned than unburned plots (P>0.04); otherwise, there were no significant differences in cover between treatments. Many germinants on burned plots did not survive. Postfire sprouting of top-killed red elderberry was observed but not quantified [111]. See the Research Project Summary of the Clearwater study for further details on the postfire response of red elderberry and 33 other plant species.

Mean percent cover (and frequency) of red elderberry before and after clearcutting and slash burning on the Clearwater Forest District of British Columbia [111]
Forest District Before logging and fire Postfire year
1 2 3 5 10
spring fire 0 (0) 1 (70) 4 (75) 1.7 (80) 2.4 (65) 1.2 (60)
fall fire 0 (0) 0.1 (53) 3.7 (47) 1.3 (53) 1 (40) 0.7 (27)
unburned 0.4 (7) 1.4 (14) 1.4 (7) 2.1 (7) 0.7 (7) 0.4 (7)

The MacKenzie and Headwaters sites were burned the summer following winter cutting. Over 10 years, red elderberry decreased slightly in cover but increased greatly in frequency over prefire values on the MacKenzie site, which was in a hybrid spruce/devil's club (P. glauca × P. engelmannii/Oplopanax horridus) forest [108]. Prefire abundance was not measured on the Headwaters site, a subalpine fir-hybrid spruce/big huckleberry forest. Red elderberry cover increased over 3 postfire years on the Headwaters site. Frequency remained stable over that period, but had decreased by postfire year 10 compared to postfire years 1 to 5 [109]. See the Research Paper of the MacKenzie study for further details on the postfire responses of red elderberry, other vascular plant species, and bryophytes.

Mean percent cover (and frequency) of red elderberry after clearcutting and slash burning on the MacKenzie [108] and Headwaters [109] Forest Districts of British Columbia
Forest District After logging;
before fire
Postfire year
1 2 3 5 10
MacKenzie 0.50 (17) 7.17 (100) 16.33 (100) 3.17 (100) 0.78 (83) 0.38 (100)
Headwaters not available 0.08 (83.3) 1.27 (83.3) 2.25 (83.3) 1.83 (100) 0.20 (50)

Herbicides and burning: Roberts [200] reported that red elderberry seedlings were "present but not abundant" 4 months after August application of picloram followed by a September prescribed fire in a red alder community on the Coast Ranges of Oregon. Red elderberry sprouting also occurred, with means of 13 inches (32 cm) for clump height, 10 inches (26 cm) for clump diameter, and 6 stems/clump for clump density at postfire month 3.5 (n=4) [200]. In the Prince George Forest District of east-central British Columbia, red elderberry established from soil-stored seed after winter clearcutting, August slash burning, and May replanting of a subalpine fir/devil's club forest. On plots where fire exposed mineral soil, red elderberry increased in the first 3 postfire years, then declined in postfire years 5 and 10. On plots where some litter and duff remained after burning, cover and frequency peaked in postfire year 1. Across postfire years 1 through 10, red elderberry was less frequent on mineral soil than on forest floor plots [107].

Mean percent cover (and frequency) of red elderberry after clearcutting and slash burning in British Columbia [107]
Substrate Prefire Postfire year
1 2 3 5 10
Mineral soil (n=19 plots) not applicable 2.90 (47.37) 7.01 (63.16) 3.21 (47.37) 1.59 (47.37) 1.70 (36.84)
Forest floor (n=128 plots) 0.09 (1.56) 15.07 (89.84) 10.70 (87.50) 3.68 (71.88) 1.50 (60.16) 0.48 (39.06)

See the Research Paper of Hamilton's [107] study for information on the responses of over 100 vascular plants, mosses, and lichens to the logging and prescribed fire treatments.

In a vine maple (Acer circinatum)-salmonberry shrubfield in coastal central Oregon, September herbicide (glyphosate) and October prescribed fire treatments reduced red elderberry cover over pretreatment levels in the short term, while September use of herbicide alone increased red elderberry cover over pretreatment levels. Red elderberries not killed by the treatments sprouted from their root crowns. Treatments were undertaken to convert the shrubfield to a conifer plantation [142].

Mean red elderberry cover (%) before and after treatments in an Oregon shrubfield [142]
Treatment Pretreatment Posttreatment year 1
Herbicide and fire 20 5
Herbicide 15 20

In a study on the Siuslaw National Forest in coastal Oregon, red elderberries on north-facing slopes recovered after combined tree harvest, spray-and-burn, and repeat spray treatments, while red elderberries on south-facing slopes were apparently favored by cutting and/or fire but killed by the second spraying. Study sites were on a Douglas-fir plantation that had been clearcut in late winter and early spring, sprayed with 2,4-D and 2,4,5-T in July, broadcast burned in September, replanted to Douglas-fir in February, and resprayed with 2,4-D and 2,4,5-T three years after the broadcast burn [210].

Mean red elderberry cover (%) after clearcutting, herbicide spraying, and burning on the Siuslaw National Forest, Oregon [210]
  North aspect South aspect
Postspray month 1 (1 month prefire) 0.05 0
Postfire year 1 0.01 0.02
Postfire year 3 1.00 0.07
Postfire year 4; 1 year after 2nd spraying 0.06 0

Red elderberry may dominate early postfire vegetation on some sites. In a chronosequence study in north-central Idaho, red elderberry was among the most abundant shrub species on logged and broadcast-burned grand fir/Oregon boxwood (Paxistima myrsinites) sites on 1-, 3-, and 8-year-old burned sites. It was 1 of 5 or 6 shrubs showing largest canopy volumes on 3- and 8-year-old burns. Red elderberry regenerated primarily by sprouting. Postfire stands dominated by red elderberry were on steep, northwest-facing slopes at the highest-elevation (5,100-5,300 feet (1,555-1,615 m)) grand fir/Oregon boxwood sites. Red elderberry canopy volume and height across different-aged stands were [249]:

Mean canopy volume and height of red elderberry on clearcut-and-broadcast burned grand fir/Oregon boxwood sites in Idaho [249]
  Postfire year 1 Postfire year 3 Postfire year 8 Postfire year 12 Postfire year 23
Canopy volume* 0.5 1.2 2.5 0.8 not given
Height (cm) 18 53 94 94 98
*Percent plant volume in a 1 × 1 × 3 m plot.

Lyon [163] reported an increase in red elderberry density after an August prescribed fire in south-central Idaho. The site was on "less disturbed terrain" within a heavily logged Douglas-fir forest. He attributed the increase in red elderberry density to multiple stems sprouting from the root crowns of formerly single-stemmed red elderberry plants. Prefire abundance of red elderberry was not quantified, but 2 years after the fire red elderberry had the 3rd largest crown volume of 12 shrub species [163]. See Lyon's Research Paper on this study for further information on the fire and the postfire responses of 64 plant species.

Mean red elderberry density and crown volume after prescribed fire in Idaho [163]
  Postfire year 1 Postfire year 2
Density (plants/1,000 feet²) 0.02 0.74
Crown volume (feet³) 0.3 27.9

Browsing: Protection from browsing will likely increase red elderberry postfire abundance on burned areas with large ungulate populations. In a sugar maple-American beech forest in the Adirondack Mountains of New York, red elderberry showed rapid growth after prescribed fall burning and postfire exclusion of white-tailed deer. Five years after treatments, red elderberry had attained heights up to 10 feet (3 m) in exclosures. On burned plots where white-tailed deer were not excluded, red elderberry was 3 feet (1 m) tall. White-tailed deer density was approximately 27 individuals/mile² in the area [19].

Frequent repeated fire: Limited studies suggest that repeated fire generally favors red elderberry [23,187], although some report red elderberry decreases after repeated fires [25,26,208]. Repeated fires generally promote red elderberry and other sprouting shrub species over conifers and fire-sensitive shrubs such as Oregon boxwood. On the Tillamook Burn of northwestern Oregon, red elderberry was not present on unburned plots but had 2% frequency on burned plots. At the time of the study, the burned plots had experienced 3 stand-replacing wildfires in 12 years. Overall, sprouting shrub species were more common in burned than unburned plots [187]. A series of prescribed fires to increase moose browse on the Chugach National Forest of southeastern Alaska slightly reduced red elderberry cover below prefire levels. Fifteen to 19 years after the fires, mean postfire coverage of red elderberry on 3 quaking aspen-balsam poplar (Populus tremuloides-P. balsamifera subsp. trichocarpa) sites was 3% compared to prefire coverage of 5% [23].

Severe fire: Based on limited studies, red elderberry shows no clear pattern of response to severe fires. Red elderberry sprouted "prolifically" the year after an explosion in a gasoline pipeline ignited a "severe" wildfire in a black cottonwood-red alder forest on Whatcom Creek, Washington [86]. It showed variable responses after severe fires in northeastern Oregon. Reestablishment was slow after a wildfire in a grand fir/queencup beadlily association near the John Day River. Red elderberry was not present on severely burned plots 1 year after the fire. It increased to 1% cover by postfire year 5. A severe wildfire near Twin Lakes, however, apparently promoted red elderberry. The fire burned through a plot established in a subalpine fir/Carolina bugbane (Trautvetteria caroliniensis) forest a year before. Red elderberry cover was 1% before the fire and 5% in postfire year 1 [133].

Late postfire succession: Since red elderberry tolerates shade (see Successional status), it may persist in late-seral postfire succession. Red elderberry was "occasionally found" on a 55-year-old burn in Yellowstone National Park [224]. In a chronosequence study in subboreal hybrid spruce/devil's club forests of British Columbia, red elderberry was a common species on 14-, 50- to 80-, and 140+-year-old burns [75].

FUELS AND FIRE REGIMES:
Fuels: Since red elderberry is a minor component of most plant communities [30,232], it does not contribute substantially to fuel loads on most sites (for example, [27,229]).

In coastal British Columbia, red elderberry is an indicator of "water-receiving" sites that have rapid decomposition of forest floor materials on burned or cutover areas [150].

Biomass and stand structure analyses: Fierke and Kauffman [83] provide equations to predict aboveground biomass of red elderberry and other woody species in black cottonwood riparian forests of the Pacific Northwest. Hanley and others [112] provide measures of leaf, twig, and stem biomass of red elderberry and other vascular plant species in red alder-western hemlock stands in southeastern Alaska. Tremblay and Larocque [229] provide measures of seasonal changes in biomass of red elderberry, other woody species, and mosses in a balsam fir stand in southern Quebec.

Gemborys [93] provides structural analyses of old-growth mixed hardwood-conifer stands on the White Mountain National Forest, New Hampshire, including densities and importance values of shrubs and basal areas, densities, and importance values of overstory trees. Red elderberry is a shrub component in some of the stands [93].

Fire regimes: Red elderberry is subject to wide variations in fire regimes across its broad geographic distribution. Since it is infrequent on most sites, fire studies to date (2008) have not determined which fire regimes favor red elderberry. Given red elderberry's position in succession—which also varies but tends toward early seral stages and open canopies—red elderberry is likely to be most abundant on sites where fire or other disturbance creates or maintains an open canopy.

Many plant communities with red elderberry historically experienced short return-interval, low-severity understory fires at intervals of 20 years or less. Examples of such communities include pine-oak (Pinus-Quercus spp.) forests and woodlands of the Appalachian Mountains [158,171], jack pine woodlands in the Great Lakes [154], and ponderosa pine (Pinus ponderosa) forests and woodlands of the West [13,222].

Many other plant communities where red elderberry is a characteristic species had mixed-severity fire regimes. Sierra lodgepole pine/mountain hemlock (P. contorta var. murrayana/Tsuga mertensiana) and Sierra lodgepole pine-western white pine (P. monticola) forests, for example, historically experienced small (5 acres (0.4 ha)), low-severity surface fires at intervals of 9 or more years [99] but also had larger, mixed surface-and-crown and crown fires at intervals of 50 or more years [99,153]. Some Rocky Mountain lodgepole pine (P. contorta var. latifolia) forests also had mixed-severity fire regimes, although many experienced mostly stand-replacement fires [1]. In a study in the northern Rocky Mountains, Arno [12] found fire was more frequent and less severe in Rocky Mountain lodgepole pine forests in areas having dry summers. Douglas-fir-western hemlock and Douglas-fir-Sitka spruce communities of the Pacific Northwest also experienced surface, surface-and-crown, and crown fires at varying intervals. In general, size and severity of fires in Douglas-fir communities historically tended to decrease, while fire frequency increased, southward from western Washington to northern California [182]. Mixed-hardwood forests of the Northeast also historically experienced some mixed-severity fires [156].

Red elderberry also occurs in plant communities that historically had mostly stand-replacement fires of various return intervals. Many spruce-fir (Picea-Abies spp.) [87,155], mixed hardwood-spruce [157], and hardwood [156] forests of the East may have gone centuries between stand-replacement fires, while relatively moist Rocky Mountain lodgepole pine forests historically experienced mostly stand-replacement fires at moderate intervals (60-80 years), with some low-severity surface fires [2,3,67]. Montane chaparral communities of the Sierra Nevada with a red elderberry component were (and are) maintained by frequent- to moderate-interval, stand-replacing fires [99]. Gap succession was probably more important than fire in maintaining red elderberry in spruce, hardwood, and mixed-wood communities that had long return-interval, stand-replacement fires.

The Fire Regime Table summarizes characteristics of fire regimes for vegetation communities in which red elderberry may occur. Follow the links in the table to documents that provide more detailed information on these fire regimes.

FIRE MANAGEMENT CONSIDERATIONS:
Prescribed and wildfires may have little effect on red elderberry abundance in many plant communities. Red elderberry may show postfire increases after either infrequent or very frequent fire. By removing the canopy, infrequent, stand-replacement fires provide openings for red elderberry establishment in plant communities that had been in the late-seral, closed canopy stage for decades or centuries. Frequent stand-replacement fire may help maintain brushfields of red elderberry, red alder, and other sprouting shrubs at the expense of conifers. Rhizomatous populations of red elderberry are probably more likely to form thickets after fire than nonrhizomatous populations. Because rhizomatous habit of red elderberry varies among populations, managers wanting to predict postfire structure of red elderberry may want to sample underground red elderberry growth on the proposed burn site.

Fire-maintained shrubfields with red elderberry are important habitat for grizzly bears. In the Selkirk Mountains of northern Idaho, grizzly bears used a mixed-shrub burn with a large red elderberry component more than expected in summer, based on the shrubfield's availability (P<0.1). One grizzly bear spent nearly 100% of her time in the burn, feeding on red elderberry and huckleberry (Vaccinium spp.) fruits [10].

MANAGEMENT CONSIDERATIONS

SPECIES: Sambucus racemosa
 

Photo used with permission of Creative Commons.

 
IMPORTANCE TO WILDLIFE AND LIVESTOCK:
Wildlife: Numerous frugivorous birds eat red elderberry fruits [35,70,101,120,170,219,236]. In the Northeast, at least 50 passerine and 6 game bird species consume the fruits [245]. Red elderberry fruits are also important in the diets of game and nongame birds in the West. An Idaho study found red elderberry fruits were a major summer diet item of blue grouse in Douglas-fir habitats [211]. A study across western Oregon found red elderberry fruits were the main summer diet item of band-tailed pigeons [130].

Mammals, including eastern fox squirrels, white-footed mice [120], other rodents [170], northern raccoons [245], American black bears [228,245], brown bears [220,228], and grizzly bears [10,39,64,105], also consume red elderberry fruits. Additionally, grizzly bears consume red elderberry foliage and roots [10,39].

Browsing ungulates consume red elderberry foliage [148,151,169,170,245], although red elderberry browse is not preferred on all sites. The browse contains cyanide [21], which is bitter, so red elderberry use may be light in areas where more palatable forage is available [122,127]. Use may be heavy, however, in areas with large white-tailed deer populations [8,245]. In winter, browsing ungulates consume red elderberry bark and buds [102]; although even then, browsing may be light if other shrubs are available [68]. Red elderberry provided minor winter forage for moose on Isle Royale, Michigan [9] and was not preferred as summer browse [21]. In contrast, by the Flathead River of Montana red elderberry was "easily the most palatable browse species on the range". Red elderberry was not abundant, and with 70% utilization by elk, it was apparently in decline [90]. Studies in coastal Alaska and the West coast found Roosevelt elk consumption of red elderberry peaked in fall (16% of total diet) and was least in spring (1% of diet) (review by [131]).

Common porcupines, mice [52,102], and snowshoe hares [68] consume red elderberry buds and bark in winter.

Various fly species consume red elderberry pollen [138]. Red elderberry and Mexican elderberry (Sambucus mexicana) are obligate hosts [223] of the federally threatened [234] valley elderberry longhorn beetle, which is endemic to the Central Valley of California [223].

Livestock: Domestic goats, sheep, and cattle browse red elderberry, most often in summer. Livestock generally prefer browsing the typical variety of red elderberry over other Sambucus taxa. Range cattle and domestic sheep on the Uinta National Forest, Utah, browsed red elderberry from late August to early September. Cattle utilization ranged from 40% to 70%; domestic sheep utilization ranged from 80% to 90% [52].

Palatability: Red elderberry browse is generally palatable to elk, deer, mountain goats, and bears [151]. Palatability of red elderberry browse increases after frost [232] and probably varies with relative cyanide content of individual plants. For deer, ratings for red elderberry browse range from moderately [180] to highly palatable [205]. Red elderberry was moderately palatable to white-tailed deer in a Wisconsin feeding trial [59]. A 4-year study in Washington found red elderberry made up 1% of the mule deer diet; most stomach samples were collected in October [31]. Sampson and Jesperson [205] reported mule deer in California consuming red elderberry foliage "with considerable relish".

Palatability of red elderberry for livestock ranges from poor to good [73]. In California, red elderberry is rated as moderately to highly palatable to domestic sheep and goats, fairly palatable to cattle, and unpalatable to horses [205]. Rocky Mountain elderberry is rated unpalatable to moderately palatable to cattle and palatable to domestic sheep [232].

Nutritional value: Although not preferred, red elderberry browse is highly nutritious. Analysis of summer browse species on Isle Royale found red elderberry had the highest mean nutrient value (crude protein + mineral content) of 28 species. However, moose browsed it in only trace amounts, less than predicted based on its nutritional content (P<0.05). The authors suggested that toxic levels of cyanide in the red elderberry browse limited moose selection [21]. See Einarsen [76] for seasonal variation in protein content of red elderberry browse. His samples were collected 6 years after one of the reburns on Tillamook Burn in Oregon [76].

Red elderberry fruits are high in carbohydrates [189,235], fat [235], and magnesium relative to most associated, fleshy-fruited shrubs of the Pacific Northwest. Norton and others [189] provide nutritional analyses of the fleshy fruits of red elderberry and other shrubs in the Pacific Northwest. See Usui and others [235] for nutritional contents of red elderberry fruits from northern Ontario.

Cover value: Red elderberries provide wildlife habitat and cover. Seral shrubfields with a red elderberry component are important grizzly bear habitat [10,15]. Sitka alder-Rocky Mountain elderberry shrubfields in the Selway-Bitterroot Wilderness of Idaho and Montana, where grizzly bears were extirpated as of 2008, have been identified as good potential grizzly bear habitat [39]. Passerines and other birds use red elderberry for nesting [102,179] and perching [120,179]. Red elderberries on streambanks provide shade cover for fish [129].

VALUE FOR REHABILITATION OF DISTURBED SITES:
Red elderberry is used in revegetation, erosion control, and wildlife plantings [43,91,120,128,179,193]. It may be relatively tolerant of heavy metal contamination; red elderberry was among a few relict shrub species present on acidic copper and nickel mining and smelting sites in Ontario [242].

Red elderberry is easily started from cuttings and is often grown from seed. See these sources for propagation information: [29,74,115,123,179,245]. Commercial sources of red elderberry are available [185].

OTHER USES:
Red elderberry fruits are used to make pies, jelly, and wine [7,80,236]. The fruits contain anthocyanins [14,188], which have antioxidant properties [14,188]. The fruits are sour, however, and people do not usually eat them raw [116,141,150]. The fruits and/or seeds may cause diarrhea and vomiting in humans [116,126,236], especially if the fruits are not fully ripe. Other parts of the plant may be poisonous to humans year-round [7].

Native Americans used red elderberry fruits as food [42,80,198,230,250] and extracts from the roots and/or bark as an emetic or purgative ([80,198,230,231], review by [172]) and to treat colds [134,198]. Native Americans also used bark extracts as a gynecological medicine and to treat influenza, fever, and tuberculosis [134,231]. The stems were used to make toys. The Dena'ina made popguns from the hollow stems, using a shelf fungus (Polyporus betulinus) for ammunition [134]. The Kwakiutl of British Columbia made toy blowguns from red elderberry stems [230].

Red elderberry is planted as an ornamental [232].

Extracts from red elderberry roots exhibited antiviral activity against bovine respiratory virus in the laboratory [172].

OTHER MANAGEMENT CONSIDERATIONS:
Red elderberry does not tolerate heavy browsing ([8], review by [102]). In Michigan, it has declined where white-tailed deer populations are high [8].

Red elderberry is a bioindicator of ozone pollution. It progresses from leaf stippling to bleaching with increasing exposure to ozone [40,168].

Control: Since red elderberry is a valuable and somewhat rare wildlife plant, control is probably not needed on most sites. Worley and Nixon [245] stated that red elderberry "seldom needs to be killed" for wildlife management purposes. Red elderberry does not greatly interfere with conifer seedling growth on most sites, although it is sometimes sprayed along with more common species, such as salmonberry, that are weedy on plantations. If chemical control of red elderberry is desired, its early leaf-out (see Seasonal development) allows for spraying early in the growing season, when conifers are less susceptible to herbicide damage [49]. Spring application of triclopyr or winter application of glyphosate caused very severe damage (90-100% dieback) to red elderberry in shrubfields in Oregon and northern Idaho [51].

APPENDIX: FIRE REGIME TABLE

SPECIES: Sambucus racemosa
Fire regime information on plant communities in which red elderberry may occur. This information is taken from the LANDFIRE Rapid Assessment Vegetation Models [160], which were developed by local experts using available literature, local data, and/or expert opinion. This table summarizes fire regime characteristics for each plant community listed. The PDF file linked from each plant community name describes the model and synthesizes the knowledge available on vegetation composition, structure, and dynamics in that community. Cells are blank where information is not available in the Rapid Assessment Vegetation Model.
Pacific Northwest California Southwest Great Basin Northern and Central Rockies
Northern Great Plains Great Lakes Northeast South-central US Southern Appalachians
Southeast        
Pacific Northwest
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northwest Grassland
Alpine and subalpine meadows and grasslands Replacement 68% 350 200 500
Mixed 32% 750 500 >1,000
Northwest Woodland
Oregon white oak-ponderosa pine Replacement 16% 125 100 300
Mixed 2% 900 50  
Surface or low 81% 25 5 30
Ponderosa pine Replacement 5% 200    
Mixed 17% 60    
Surface or low 78% 13    
Oregon white oak Replacement 3% 275    
Mixed 19% 50    
Surface or low 78% 12.5    
Subalpine woodland Replacement 21% 300 200 400
Mixed 79% 80 35 120
Northwest Forested
Sitka spruce-western hemlock Replacement 100% 700 300 >1,000
Douglas-fir (Willamette Valley foothills) Replacement 18% 150 100 400
Mixed 29% 90 40 150
Surface or low 53% 50 20 80
Oregon coastal tanoak Replacement 10% 250    
Mixed 90% 28 15 40
Dry ponderosa pine (mesic) Replacement 5% 125    
Mixed 13% 50    
Surface or low 82% 8    
Douglas-fir-western hemlock (dry mesic) Replacement 25% 300 250 500
Mixed 75% 100 50 150
Douglas-fir-western hemlock (wet mesic) Replacement 71% 400    
Mixed 29% >1,000    
Mixed conifer (southwestern Oregon) Replacement 4% 400    
Mixed 29% 50    
Surface or low 67% 22    
California mixed evergreen (northern California) Replacement 6% 150 100 200
Mixed 29% 33 15 50
Surface or low 64% 15 5 30
Mountain hemlock Replacement 93% 750 500 >1,000
Mixed 7% >1,000    
Pacific silver fir (low elevation) Replacement 46% 350 100 800
Mixed 54% 300 100 400
Pacific silver fir (high elevation) Replacement 69% 500    
Mixed 31% >1,000    
Subalpine fir Replacement 81% 185 150 300
Mixed 19% 800 500 >1,000
Mixed conifer (eastside mesic) Replacement 35% 200    
Mixed 47% 150    
Surface or low 18% 400    
Red fir Replacement 20% 400 150 400
Mixed 80% 100 80 130
Spruce-fir Replacement 84% 135 80 270
Mixed 16% 700 285 >1,000
California
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
California Grassland
California grassland Replacement 100% 2 1 3
Herbaceous wetland Replacement 70% 15    
Mixed 30% 35    
Wet mountain meadow-Lodgepole pine (subalpine) Replacement 21% 100    
Mixed 10% 200    
Surface or low 69% 30    
Alpine meadows and barrens Replacement 100% 200 200 400
California Shrubland
Coastal sage scrub Replacement 100% 50 20 150
Coastal sage scrub-coastal prairie Replacement 8% 40 8 900
Mixed 31% 10 1 900
Surface or low 62% 5 1 6
Montane chaparral Replacement 34% 95    
Mixed 66% 50    
California Woodland
California oak woodlands Replacement 8% 120    
Mixed 2% 500    
Surface or low 91% 10    
Ponderosa pine Replacement 5% 200    
Mixed 17% 60    
Surface or low 78% 13    
California Forested
California mixed evergreen Replacement 10% 140 65 700
Mixed 58% 25 10 33
Surface or low 32% 45 7  
Coast redwood Replacement 2% ≥1,000    
Surface or low 98% 20    
Mixed conifer (North Slopes) Replacement 5% 250    
Mixed 7% 200    
Surface or low 88% 15 10 40
Mixed conifer (South Slopes) Replacement 4% 200    
Mixed 16% 50    
Surface or low 80% 10    
Aspen with conifer Replacement 24% 155 50 300
Mixed 15% 240    
Surface or low 61% 60    
Jeffrey pine Replacement 9% 250    
Mixed 17% 130    
Surface or low 74% 30    
Mixed evergreen-bigcone Douglas-fir (southern coastal) Replacement 29% 250    
Mixed 71% 100    
Interior white fir (northeastern California) Replacement 47% 145    
Mixed 32% 210    
Surface or low 21% 325    
Red fir-white fir Replacement 13% 200 125 500
Mixed 36% 70    
Surface or low 51% 50 15 50
Red fir-western white pine Replacement 16% 250    
Mixed 65% 60 25 80
Surface or low 19% 200    
Sierra Nevada lodgepole pine (cold wet upper montane) Replacement 23% 150 37 764
Mixed 70% 50    
Surface or low 7% 500    
Southwest
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Southwest Grassland
Montane and subalpine grasslands with shrubs or trees Replacement 30% 70 10 100
Surface or low 70% 30    
Southwest Forested
Riparian forest with conifers Replacement 100% 435 300 550
Riparian deciduous woodland Replacement 50% 110 15 200
Mixed 20% 275 25  
Surface or low 30% 180 10  
Ponderosa pine-Douglas-fir (southern Rockies) Replacement 15% 460    
Mixed 43% 160    
Surface or low 43% 160    
Southwest mixed conifer (cool, moist with aspen) Replacement 29% 200 80 200
Mixed 35% 165 35  
Surface or low 36% 160 10  
Aspen with spruce-fir Replacement 38% 75 40 90
Mixed 38% 75 40  
Surface or low 23% 125 30 250
Stable aspen without conifers Replacement 81% 150 50 300
Surface or low 19% 650 600 >1,000
Lodgepole pine (Central Rocky Mountains, infrequent fire) Replacement 82% 300 250 500
Surface or low 18% >1,000 >1,000 >1,000
Spruce-fir Replacement 96% 210 150  
Mixed 4% >1,000 35 >1,000
Great Basin
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Great Basin Grassland
Great Basin grassland Replacement 33% 75 40 110
Mixed 67% 37 20 54
Mountain meadow (mesic to dry) Replacement 66% 31 15 45
Mixed 34% 59 30 90
Great Basin Shrubland
Montane chaparral Replacement 37% 93    
Mixed 63% 54    
Great Basin Woodland
Ponderosa pine Replacement 5% 200    
Mixed 17% 60    
Surface or low 78% 13    
Great Basin Forested
Interior ponderosa pine Replacement 5% 161   800
Mixed 10% 80 50 80
Surface or low 86% 9 8 10
Ponderosa pine-Douglas-fir Replacement 10% 250   >1,000
Mixed 51% 50 50 130
Surface or low 39% 65 15  
Aspen with conifer (low to midelevation) Replacement 53% 61 20  
Mixed 24% 137 10  
Surface or low 23% 143 10  
Douglas-fir (warm mesic interior) Replacement 28% 170 80 400
Mixed 72% 65 50 250
Aspen with conifer (high elevation) Replacement 47% 76 40  
Mixed 18% 196 10  
Surface or low 35% 100 10  
Stable aspen-cottonwood, no conifers Replacement 31% 96 50 300
Surface or low 69% 44 20 60
Spruce-fir-pine (subalpine) Replacement 98% 217 75 300
Mixed 2% >1,000    
Aspen with spruce-fir Replacement 38% 75 40 90
Mixed 38% 75 40  
Surface or low 23% 125 30 250
Stable aspen without conifers Replacement 81% 150 50 300
Surface or low 19% 650 600 >1,000
Northern and Central Rockies
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northern and Central Rockies Grassland
Northern prairie grassland Replacement 55% 22 2 40
Mixed 45% 27 10 50
Mountain grassland Replacement 60% 20 10  
Mixed 40% 30    
Northern and Central Rockies Shrubland
Riparian (Wyoming)
Mixed 100% 100 25 500
Mountain shrub, nonsagebrush Replacement 80% 100 20 150
Mixed 20% 400    
Northern and Central Rockies Woodland
Ancient juniper Replacement 100% 750 200 >1,000
Northern and Central Rockies Forested
Ponderosa pine (Northern Great Plains) Replacement 5% 300    
Mixed 20% 75    
Surface or low 75% 20 10 40
Ponderosa pine (Northern and Central Rockies) Replacement 4% 300 100 >1,000
Mixed 19% 60 50 200
Surface or low 77% 15 3 30
Ponderosa pine (Black Hills, low elevation) Replacement 7% 300 200 400
Mixed 21% 100 50 400
Surface or low 71% 30 5 50
Ponderosa pine (Black Hills, high elevation) Replacement 12% 300    
Mixed 18% 200    
Surface or low 71% 50    
Ponderosa pine-Douglas-fir Replacement 10% 250   >1,000
Mixed 51% 50 50 130
Surface or low 39% 65 15  
Western redcedar Replacement 87% 385 75 >1,000
Mixed 13% >1,000 25  
Douglas-fir (warm mesic interior) Replacement 28% 170 80 400
Mixed 72% 65 50 250
Douglas-fir (cold) Replacement 31% 145 75 250
Mixed 69% 65 35 150
Grand fir-Douglas-fir-western larch mix Replacement 29% 150 100 200
Mixed 71% 60 3 75
Mixed conifer-upland western redcedar-western hemlock Replacement 67% 225 150 300
Mixed 33% 450 35 500
Western larch-lodgepole pine-Douglas-fir Replacement 33% 200 50 250
Mixed 67% 100 20 140
Grand fir-lodgepole pine-larch-Douglas-fir Replacement 31% 220 50 250
Mixed 69% 100 35 150
Persistent lodgepole pine Replacement 89% 450 300 600
Mixed 11% >1,000    
Lower subalpine lodgepole pine Replacement 73% 170 50 200
Mixed 27% 450 40 500
Lower subalpine (Wyoming and Central Rockies) Replacement 100% 175 30 300
Upper subalpine spruce-fir (Central Rockies) Replacement 100% 300 100 600
Northern Great Plains
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northern Plains Woodland
Oak woodland Replacement 2% 450    
Surface or low 98% 7.5
Northern Great Plains wooded draws and ravines Replacement 38% 45 30 100
Mixed 18% 94    
Surface or low 43% 40 10  
Great Plains floodplain Replacement 100% 500    
Great Lakes
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Great Lakes Woodland
Great Lakes pine barrens Replacement 8% 41 10 80
Mixed 9% 36 10 80
Surface or low 83% 4 1 20
Jack pine-open lands (frequent fire-return interval) Replacement 83% 26 10 100
Mixed 17% 125 10  
Northern oak savanna Replacement 4% 110 50 500
Mixed 9% 50 15 150
Surface or low 87% 5 1 20
Great Lakes Forested
Northern hardwood maple-beech-eastern hemlock Replacement 60% >1,000    
Mixed 40% >1,000    
Conifer lowland (embedded in fire-prone system) Replacement 45% 120 90 220
Mixed 55% 100    
Conifer lowland (embedded in fire-resistant ecosystem) Replacement 36% 540 220 >1,000
Mixed 64% 300    
Great Lakes floodplain forest
Mixed 7% 833    
Surface or low 93% 61    
Great Lakes spruce-fir Replacement 100% 85 50 200
Minnesota spruce-fir (adjacent to Lake Superior and Drift and Lake Plain) Replacement 21% 300    
Surface or low 79% 80    
Great Lakes pine forest, jack pine Replacement 67% 50    
Mixed 23% 143    
Surface or low 10% 333
Maple-basswood Replacement 33% >1,000    
Surface or low 67% 500    
Maple-basswood mesic hardwood forest (Great Lakes) Replacement 100% >1,000 >1,000 >1,000
Maple-basswood-oak-aspen Replacement 4% 769    
Mixed 7% 476    
Surface or low 89% 35    
Northern hardwood-eastern hemlock forest (Great Lakes) Replacement 99% >1,000    
Oak-hickory Replacement 13% 66 1  
Mixed 11% 77 5  
Surface or low 76% 11 2 25
Pine-oak Replacement 19% 357    
Surface or low 81% 85    
Red pine-eastern white pine (frequent fire) Replacement 38% 56    
Mixed 36% 60    
Surface or low 26% 84    
Red pine-eastern white pine (less frequent fire) Replacement 30% 166    
Mixed 47% 105    
Surface or low 23% 220    
Great Lakes pine forest, eastern white pine-eastern hemlock (frequent fire) Replacement 52% 260    
Mixed 12% >1,000    
Surface or low 35% 385    
Eastern white pine-eastern hemlock Replacement 54% 370    
Mixed 12% >1,000    
Surface or low 34% 588    
Northeast
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northeast Woodland
Eastern woodland mosaic Replacement 2% 200 100 300
Mixed 9% 40 20 60
Surface or low 89% 4 1 7
Rocky outcrop pine (Northeast) Replacement 16% 128    
Mixed 32% 65    
Surface or low 52% 40    
Pine barrens Replacement 10% 78    
Mixed 25% 32    
Surface or low 65% 12    
Northeast Forested
Northern hardwoods (Northeast) Replacement 39% >1,000    
Mixed 61% 650    
Eastern white pine-northern hardwoods Replacement 72% 475    
Surface or low 28% >1,000    
Northern hardwoods-eastern hemlock Replacement 50% >1,000    
Surface or low 50% >1,000    
Northern hardwoods-spruce Replacement 100% >1,000 400 >1,000
Appalachian oak forest (dry-mesic) Replacement 2% 625 500 >1,000
Mixed 6% 250 200 500
Surface or low 92% 15 7 26
Beech-maple Replacement 100% >1,000    
Northeast spruce-fir forest Replacement 100% 265 150 300
Southeastern red spruce-Fraser fir Replacement 100% 500 300 >1,000
South-central US
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
South-central US Woodland
Oak-hickory savanna Replacement 1% 227    
Surface or low 99% 3.2    
Oak woodland-shrubland-grassland mosaic Replacement 11% 50    
Mixed 56% 10    
Surface or low 33% 17    
Interior Highlands oak-hickory-pine Replacement 3% 150 100 300
Surface or low 97% 4 2 10
South-central US Forested
Interior Highlands dry-mesic forest and woodland Replacement 7% 250 50 300
Mixed 18% 90 20 150
Surface or low 75% 22 5 35
Southern Appalachians
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Southern Appalachians Grassland
Bluestem-oak barrens Replacement 46% 15    
Mixed 10% 69    
Surface or low 44% 16    
Eastern prairie-woodland mosaic Replacement 50% 10    
Mixed 1% 900    
Surface or low 50% 10    
Southern Appalachians Woodland
Appalachian shortleaf pine Replacement 4% 125    
Mixed 4% 155    
Surface or low 92% 6    
Table Mountain-pitch pine Replacement 5% 100    
Mixed 3% 160    
Surface or low 92% 5    
Oak-ash woodland Replacement 23% 119    
Mixed 28% 95    
Surface or low 49% 55    
Southern Appalachians Forested
Bottomland hardwood forest Replacement 25% 435 200 >1,000
Mixed 24% 455 150 500
Surface or low 51% 210 50 250
Mixed mesophytic hardwood Replacement 11% 665    
Mixed 10% 715    
Surface or low 79% 90    
Appalachian oak-hickory-pine Replacement 3% 180 30 500
Mixed 8% 65 15 150
Surface or low 89% 6 3 10
Eastern hemlock-eastern white pine-hardwood Replacement 17% >1,000 500 >1,000
Surface or low 83% 210 100 >1,000
Appalachian Virginia pine Replacement 20% 110 25 125
Mixed 15% 145    
Surface or low 64% 35 10 40
Appalachian oak forest (dry-mesic) Replacement 6% 220    
Mixed 15% 90    
Surface or low 79% 17    
Southern Appalachian high-elevation forest Replacement 59% 525    
Mixed 41% 770    
Southeast
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Southeast Woodland
Longleaf pine/bluestem Replacement 3% 130    
Surface or low 97% 4 1 5
Longleaf pine (mesic uplands) Replacement 3% 110 40 200
Surface or low 97% 3 1 5
Atlantic wet pine savanna Replacement 4% 100    
Mixed 2% 175    
Surface or low 94% 4     
Southeast Forested
Atlantic white-cedar forest Replacement 34% 200 25 350
Mixed 8% 900 20 900
Surface or low 59% 115 10 500
*Fire Severities
Replacement: Any fire that causes greater than 75% top removal of a vegetation-fuel type, resulting in general replacement of existing vegetation; may or may not cause a lethal effect on the plants.
Mixed: Any fire burning more than 5% of an area that does not qualify as a replacement, surface, or low-severity fire; includes mosaic and other fires that are intermediate in effects.
Surface or low: Any fire that causes less than 25% upper layer replacement and/or removal in a vegetation-fuel class but burns 5% or more of the area [113,159].

Sambucus racemosa: REFERENCES


1. Agee, James K. 1993. Fire ecology of Pacific Northwest forests. Washington, DC: Island Press. 493 p. [22247]
2. Agee, James K. 1994. Fire and weather disturbances in terrestrial ecosystems of the eastern Cascades. Gen. Tech. Rep. PNW-GTR-320. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 52 p. [Hessburg, Paul F., tech. ed. Eastside forest ecosystem health assessment. Vol. 3: assessment]. [23656]
3. Agee, James K. 1998. Fire and pine ecosystems. In: Richardson, David M., ed. Ecology and biogeography of Pinus. Cambridge, United Kingdom: The Press Syndicate of the University of Cambridge: 193-218. [37704]
4. Ahlgren, Clifford E. 1979. Buried seed in the forest floor of the Boundary Waters Canoe Area. Minnesota Forestry Research Note No. 271. St. Paul, MN: University of Minnesota, College of Forestry. 4 p. [3459]
5. Alaback, Paul B.; Herman, F. R. 1988. Long-term response of understory vegetation to stand density in Picea-Tsuga forests. Canadian Journal of Forest Research. 18: 1522-1530. [6227]
6. Alaback, Paul B.; Tappeiner, John C., II. 1991. Response of western hemlock (Tsuga heterophylla) and early huckleberry (Vaccinium ovalifolium) seedlings to forest windthrow. Canadian Journal of Forest Research. 21: 534-539. [15051]
7. Alderman, DeForest C. 1979. Native edible fruits, nuts, vegetables, herbs, spices, and grasses of California: II. Small or bushy fruits. Leaflet 2278. Berkeley, CA: University of California, Division of Agricultural Sciences, Cooperative Extension. 26 p. [67652]
8. Aldous, Shaler E. 1952. Deer browse clipping study in the Lake States Region. Journal of Wildlife Management. 16(4): 401-409. [6826]
9. Aldous, Shaler E.; Krefting, Laurits W. 1946. The present status of moose on Isle Royale. Transactions, 11th North American Wildlife Conference. 11: 296-308. [17042]
10. Almack, Jon. 1986. Grizzly bear habitat use, food habits, and movements in the Selkirk Mountains, northern Idaho. In: Contreras, Glen P.; Evans, Keith E., compilers. Proceedings--grizzly bear habitat symposium; 1985 April 30 - May 2; Missoula, MT. Gen. Tech. Rep. INT-207. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 150-157. [10815]
11. Archambault, Louis; Morissette, Jacques; Bernier-Cardou, Michele. 1998. Forest succession over a 20-year period following clearcutting in balsam fir--yellow birch ecosystems of eastern Quebec, Canada. Forest Ecology and Management. 102(1): 61-74. [28451]
12. Arno, Stephen F. 1980. Forest fire history in the Northern Rockies. Journal of Forestry. 78(8): 460-465. [11990]
13. 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]
14. Bagchi, D.; Sen, C. K.; Bagchi, M.; Atalay, M. 2004. Anti-angiogenic, antioxidant, and anti-carcinogenic properties of a novel anthocyanin-rich berry extract formula. Biochemistry. 69(1): 75-80. [68988]
15. Banner, Allen; Pojar, Jim; Trowbridge, Rick; Hamilton, Anthony. 1986. Grizzly bear habitat in the Kimsquit River Valley, coastal British Columbia: classification, description, and mapping. In: Contreras, Glen P.; Evans, Keith E., compilers. Proceedings--grizzly bear habitat symposium; 1985 April 30 - May 2; Missoula, MT. Gen. Tech. Rep. INT-207. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 36-49. [10810]
16. Barbour, M. G.; Fernau, R. F.; Benayas, J. M. Rey; Jurjavcic, E. B.; Royce, E. B. 1998. Tree regeneration following clearcut logging in red fir forests of California. Forest Ecology and Management. 104: 101-111. [29227]
17. Baskin, Carol C.; Baskin, Jerry M. 2001. Seeds: ecology, biogeography, and evolution of dormancy and germination. San Diego, CA: Academic Press. 666 p. [60775]
18. Batchelor, Ron; Erwin, Mike; Martinka, Robert; McIntosh, Don; Pfister, Robert; Schneegas, Edward; Taylor, Jack; Walther, Kit. 1982. A taxonomic classification system for Montana riparian vegetation types: An interagency approach to classifying Montana's riparian ecosystems. Bozeman, MT: Montana State Rural Areas Development Committee, Wildlife Subcommittee, Riparian Program Team. 13 p. [31151]
19. Behrend, Donald F.; Patric, Earl F. 1969. Influence of site disturbance and removal of shade on regeneration of deer browse. Journal of Wildlife Management. 33(2): 394-398. [15619]
20. Belcher, Earl. 1985. Handbook on seeds of browse-shrubs and forbs. Tech. Publ. R8-TP8. Atlanta, GA: U.S. Department of Agriculture, Forest Service, Southern Region. 246 p. In cooperation with: Association of Official Seed Analysts. [43463]
21. Belovsky, Gary E. 1981. Food plant selection by a generalist herbivore: the moose. Ecology. 62(4): 1020-1030. [64420]
22. Boles, Patrick H.; Dick-Peddie, William A. 1983. Woody riparian vegetation patterns on a segment of the Mimbres River in southwestern New Mexico. The Southwestern Naturalist. 28(1): 81-87. [65317]
23. Boucher, Tina V. 2003. Vegetation response to prescribed fire in the Kenai Mountains, Alaska. Res. Pap. PNW-RP-554. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 59 p. [48392]
24. Boucher, Tina V.; Mead, Bert R. 2006. Vegetation change and forest regeneration on the Kenai Peninsula, Alaska following a spruce beetle outbreak, 1987-2000. Forest Ecology and Management. 227(3): 233-246. [66316]
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. Bradley, Anne F.; Noste, Nonan V.; Fischer, William C. 1992. Fire ecology of forests and woodlands in Utah. Gen. Tech. Rep. INT-287. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 128 p. [18211]
27. 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]
28. Brink, V. C. 1954. Survival of plants under flood in the lower Fraser River valley, British Columbia. Ecology. 35(1): 94-95. [64483]
29. Brinkman, Kenneth A.; Johnson, Gary. [In press]. Sambucus L.--elder, [Online]. In: Bonner, Franklin T.; Nisley, Rebecca G.; Karrfait, R. P., coords. Woody plant seed manual. Agric. Handbook 727. Washington, DC: U.S. Department of Agriculture, Forest Service (Producer). Available: http://www.nsl.fs.fed.us/wpsm/Sambucus.pdff [2008, September 12]. [71055]
30. Brown, Dalton Milford. 1941. Vegetation of Roan Mountain: a phytosociological and successional study. Ecological Monographs. 11: 61-97. [23349]
31. Brown, Ellsworth R. 1961. The black-tailed deer of western Washington. Biological Bulletin No. 13. Olympia, WA: Washington State Game Commission. 124 p. [8843]
32. Brumelis, G.; Carleton, T. J. 1989. The vegetation of post-logged black spruce lowlands in central Canada. II. Understory vegetation. Journal of Applied Ecology. 26: 321-339. [7864]
33. Brundrett, Mark C.; Kendrick, Bryce. 1988. The mycorrhizal status, root anatomy, and phenology of plants in a sugar maple forest. Canadian Journal of Botany. 66(6): 1153-1173. [14483]
34. Buell, Murray F.; Wilbur, Robert L. 1948. Life-form spectra of the hardwood forests of the Itasca Park region, Minnesota. Ecology. 29(3): 352-359. [554]
35. Burns, K. C. 2003. Broad-scale reciprocity in an avian seed dispersal mutualism. Global Ecology and Biogeography. 12(5): 421-426. [69000]
36. Burns, K. C. 2005. Is there limiting similarity in the phenology of fleshy fruits? Journal of Vegetation Science. 16(6): 617-624. [68999]
37. Burns, K. C. 2007. Patterns in the assembly of an island plant community. Journal of Biogeography. 34 (5): 760-768. [68981]
38. Busing, Richard T.; Clebsch, Edward E. C.; Eagar, Christopher C.; Pauley, Eric F. 1988. Two decades of change in a Great Smoky Mountains spruce-fir forest. Bulletin of the Torrey Botanical Club. 115(1): 25-31. [4491]
39. Butterfield, Bart R.; Almack, Jon A. 1985. Evaluation of grizzly bear habitat in the Selway-Bitterroot Wilderness Area. Moscow, ID: University of Idaho, Cooperative Wildlife Research Unit. Final Report IDFG Project No. 04-78-719. 66 p. [30044]
40. Campbell, Sally; Smith, Gretchen; Temple, Pat; Pronos, John; Rochefort, Regina; Andersen, Chris. 2000. Monitoring for ozone injury in West Coast (Oregon, Washington, California) Forest in 1998. Gen. Tech. Rep. PNW-GTR-495. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 19 p. [68996]
41. Carlton, Gary C. 1988. The structure and dynamics of red alder communities in the central Coast Range of western Oregon. Corvallis, OR: Oregon State University. 173 p. Thesis. [10549]
42. Castetter, Edward F. 1935. Ethnobiological studies in the American Southwest. Biological Series No. 4: Volume 1. Albuquerque, NM: University of New Mexico. 62 p. [35938]
43. Chan, Franklin J. 1993. Response of revegetation on a severely disturbed decomposed granitic site. In: Sommarstrom, Sari, ed. Proceedings of the conference on decomposed granitic soils: problems and solutions; 1992 October 21-23. Redding, CA; University of California, Davis, University Extension: 140-151. [27532]
44. Chapman, F. C. 1947. The elderberries in wildlife conservation. Ohio Conservation Bulletin. 11(8): 20-21. [6922]
45. Clark, David Lee. 1991. The effect of fire on Yellowstone ecosystem seed banks. Bozeman, MT: Montana State University. 115 p. Thesis. [36504]
46. Clement, C. J. E. 1985. Floodplain succession on the west coast of Vancouver Island. Canadian Field-Naturalist. 99(1): 34-39. [8928]
47. Cockerell, T. D. A. 1891. Notes on the flora of high altitudes in Custer County, Colorado. Bulletin of the Torrey Botanical Club. 18(6): 167-174. [62924]
48. Coffman, Michael S.; Alyanak, Edward; Resovsky, Richard. 1980. Habitat classification system field guide: Northern Lake States region--Upper Peninsula of Michigan and northeast Wisconsin. Houghton, MI: Michigan Technological University. 112 p. [Developed by: Cooperative research on forest soils.]. [8997]
49. Cole, E. C.; Newton, M.; Youngblood, A. 1999. Regenerating white spruce, paper birch, and willow in south-central Alaska. Canadian Journal of Forest Research. 29: 993-1001. [38114]
50. Cole, Elizabeth C.; Newton, Michael; White, Diane E. 1987. Evaluation of herbicides for early season conifer release. Proceedings, Western Society of Weed Science. 40: 119-128. [38437]
51. Conard, Susan G.; Emmingham, W. H. 1983. Herbicides for shrub control on forest sites in northeastern Oregon and northern Idaho. Special Publication 5. Corvallis, OR: Oregon State University, College of Forestry, Forest Research Laboratory. 7 p. [3579]
52. Conrad, P. W.; McDonough, W. T. 1972. Growth and reproduction of red elderberry on subalpine rangeland in Utah. Northwest Science. 46(2): 140_148. [6855]
53. Cooper, William S. 1913. The climax forest of Isle Royale, Lake Superior, and its development. II. Botanical Gazette. 55(2): 115-140. [11538]
54. Cooper, William S. 1942. Vegetation of the Prince William Sound region, Alaska; with a brief excursion into post-Pleistocene climatic history. Ecological Monographs. 12(1): 1-22. [62718]
55. Cooper, William Skinner. 1923. The recent ecological history of Glacier Bay, Alaska: the present vegetation cycle. Ecology. 4(3): 223-246. [65668]
56. Cowan, Ian McTaggart. 1945. The ecological relationships of the food of the Columbian black-tailed deer, Odocoileus hemionus columbianus (Richardson), in the coast forest region of southern Vancouver Island, British Columbia. Ecological Monographs. 15(2): 110-139. [16006]
57. Cronquist, Arthur; Holmgren, Arthur H.; Holmgren, Noel H.; Reveal, James L.; Holmgren, Patricia K. 1984. Intermountain flora: Vascular plants of the Intermountain West, U.S.A. Vol. 4: Subclass Asteridae, (except Asteraceae). New York: The New York Botanical Garden. 573 p. [718]
58. Curtis, John T. 1959. Northern forests-mesic. In: Curtis, John T. The vegetation of Wisconsin. Madison, WI: The University of Wisconsin Press: 184-201. [60522]
59. Dahlberg, B. L.; Guettinger, R. C. 1956. The white-tailed deer in Wisconsin. Madison, WI: Wisconsin Conservation Department. 282 p. [71489]
60. Dansereau, Pierre. 1959. Phytogeographia Laurentiana. II. The principal plant associations of the Saint Lawrence Valley. Contributions of the Botanical Institute No. 75. Montreal, PQ: University of Montreal, Botanical Institute. 147 p. [8925]
61. Daubenmire, Rexford F. 1936. The "big woods" of Minnesota: its structure, and relation to climate, fire, and soils. Ecological Monographs. 6(2): 233-268. [2697]
62. Daubenmire, Rexford. 1981. Subalpine parks associated with snow transfer in the mountains of northern Idaho and eastern Washington. Northwest Science. 55(2): 124-135. [8273]
63. Davidson, Donald W. 1973. Vascular plants of the mature upland forests of northern New Jersey. Bulletin of the Torrey Botanical Club. 100(1): 44-55. [62587]
64. Davis, Dan; Butterfield, Bart. 1991. The Bitterroot Grizzly Bear Evaluation Area: A report to the Bitterroot Technical Review Team. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 56 p. [30041]
65. Davis, James N. 2004. Climate and terrain. In: Monsen, Stephen B.; Stevens, Richard; Shaw, Nancy L., comps. Restoring western ranges and wildlands. Gen. Tech. Rep. RMRS-GTR-136-vol-1. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 33-38. [52821]
66. Davis, John H., Jr. 1930. Vegetation of the Black Mountains of North Carolina: an ecological study. Journal of the Elisha Mitchell Scientific Society. 29: 291-318. [64613]
67. Davis, Kathleen M.; Clayton, Bruce D.; Fischer, William C. 1980. Fire ecology of Lolo National Forest habitat types. INT-79. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 77 p. [5296]
68. de Vos, Antoon. 1964. Food utilization of snowshoe hares on Mantioulin Island, Ontario. Journal of Forestry. 62: 238-244. [25071]
69. DeMeo, Tom; Martin, Jon; West, Randolph A. 1993. Forest plant association management guide: Ketchikan Area, Tongass National Forest. R10-MB-210. Juneau, AK: U.S. Department of Agriculture, Forest Service, Alaska Region. 405 p. [22498]
70. Denslow, J. S. 1987. Fruit removal rates from aggregated and isolates bushes of the red elderberry, Sambucus pubens. Canadian Journal of Botany. 65(6): 1229-1235. [6846]
71. Denslow, Julie Sloan. 1987. Fruit removal from aggregated and isolated bushes of the red elderberry Sambucus pubens. Canadian Journal of Botany. 65(6): 1229-1235. [68991]
72. DeSelm, H. R.; Boner, R. R. 1984. Understory changes in spruce-fir during the first 16-20 years following the death of fir. In: White, Peter S., ed. Southern Appalachian spruce-fir ecosystem: its biology and threats. Research/Resources Management Report SER-71. Atlanta, GA: U.S. Department of the Interior, National Park Service, Southeast Region: 51-69. [21927]
73. Dittberner, Phillip L.; Olson, Michael R. 1983. The Plant Information Network (PIN) data base: Colorado, Montana, North Dakota, Utah, and Wyoming. FWS/OBS-83/86. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service. 786 p. [806]
74. Doran, William L. 1957. Propagation of woody plants by cuttings. Experiment Station Bulletin No. 491. Amherst, MA: University of Massachusetts, College of Agriculture. 99 p. [6399]
75. Driscoll, K. G.; Arocena, J. M.; Massicotte, H. B. 1999. Post-fire soil nitrogen content and vegetation composition in sub-boreal spruce forests of British Columbia's central interior, Canada. Forest Ecology and Management. 121: 227-237. [30330]
76. Einarsen, Arthur S. 1946. Crude protein determination of deer food as an applied management technique. Transactions, 11th North American Wildlife Conference. 11: 309-312. [17031]
77. Eis, S. 1981. Effect of vegetative competition on regeneration of white spruce. Canadian Journal of Forest Research. 11: 1-8. [10104]
78. Elliott, Jack C. 1953. Composition of upland second growth hardwood stands in the tension zone of Michigan as affected by soils and man. Ecological Monographs. 23(3): 271-288. [64460]
79. Ellison, Lincoln. 1949. Establishment of vegetation on depleted subalpine range as influenced by microenvironment. Ecological Monographs. 19(2): 95-121. [66941]
80. Everett, Yvonne. 1997. A guide to selected non-timber forest products of the Hayfork Adaptive Management Area, Shasta-Trinity and Six Rivers National Forests, California. Gen. Tech. Rep. PSW-GTR-162. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station. 64 p. [28986]
81. Farrar, John Laird. 1995. Trees of the northern United States and Canada. Ames, IA: Blackwell Publishing. 502 p. [60614]
82. Ferguson, Dennis E.; Johnson, Frederic D. 1996. Classification of grand fir mosaic habitats. Gen. Tech. Rep. INT-GTR-337. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 16 p. [26770]
83. Fierke, Melissa K.; Kauffman, J. Boone. 2005. Structural dynamics of riparian forests along a black cottonwood successional gradient. Forest Ecology and Management. 215(1-3): 149-162. [55572]
84. Flaccus, Edward; Ohmann, Lewis F. 1964. Old-growth northern hardwood forests in northeastern Minnesota. Ecology. 45(3): 448-459. [49631]
85. Fleming, G. P.; Coulling, P. P.; Patterson, K. D. 2005. Terrestrial system, [Online]. In: The natural communities of Virginia: Classification of ecological community groups. Second approximation. Version 2.1. Richmond, VA: Virginia Department of Conservation and Recreation, Division of Natural Heritage (Producer). Available: http://www.dcr.virginia.gov/dnh/ncintro.htm [2005, November 3]. [60507]
86. Fonda, R. W. 2001. Postfire response of red alder, black cottonwood, and bigleaf maple to the Whatcom Creek fire, Bellingham, Washington. Northwest Science. 75(1): 25-36. [38964]
87. Foster, David R. 1985. Vegetation development following fire in Picea mariana (black spruce)-Pleurozium forests of south-eastern Labrador, Canada. Journal of Ecology. 73: 517-534. [7222]
88. Franklin, Jerry F.; Dyrness, C. T. 1973. Natural vegetation of Oregon and Washington. Gen. Tech. Rep. PNW-8. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 417 p. [961]
89. Franklin, Jerry F.; Pechanec, Anna A. 1968. Comparison of vegetation in adjacent alder, conifer, and mixed alder-conifer communities. I. Understory vegetation and stand structure. In: Trappe, J. M.; Franklin, J. F.; Tarrant, R. F.; Hansen, G. M., eds. Biology of alder: Proceedings of a symposium: 40th annual meeting of the Northwest Scientific Association; 1967 April 14-15; Pullman, WA. Portland, OR: U. S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station: 37-43. [6188]
90. Gaffney, William S. 1941. The effects of winter elk browsing, South Fork of the Flathead River, Montana. Journal of Wildlife Management. 5(4): 427-453. [5028]
91. Gardner, R. B.; Hungerford, R. D. 1975. Evaluating road design, construction, and revegetation alternatives. In: Forest residues utilization research and development program. Progress Rep. 1. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station: 41-49. [15412]
92. Geier-Hayes, Kathleen. 1991. Natural regeneration microsites for Douglas-fir in central Idaho. In: Baumgartner, David M.; Lotan, James E., compilers. Interior Douglas-fir: The species and its management: Symposium proceedings; 1991 February 27 - March 1; Spokane, WA. Pullman, WA: Washington State University, Cooperative Extension: 247-254. [18299]
93. Gemborys, Stanley R. 1996. Structure and dynamics in a virgin northern hardwood-spruce-fir forest-- The Bowl, New Hampshire. Res. Pap. NE-704. Radnor, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 15 p. [27061]
94. Gill, John D.; Healy, William M. 1974. Shrubs and vines for northeastern wildlife. Gen. Tech. Rep. NE-9. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 180 p. [6207]
95. Gleason, Henry A.; Cronquist, Arthur. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York: New York Botanical Garden. 910 p. [20329]
96. Graber, Raymond E. 1980. Pin cherry. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 17-18. [45759]
97. Graber, Raymond E.; Thompson, Donald F. 1978. Seeds in the organic layers and soil of four beech-birch-maple stands. Res. Pap. NE-401. Broomall, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 8 p. [45843]
98. Great Plains Flora Association. 1986. Flora of the Great Plains. Lawrence, KS: University Press of Kansas. 1392 p. [1603]
99. 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]
100. Gruell, George E.; Brown, James K.; Bushey, Charles L. 1986. Prescribed fire opportunities in grasslands invaded by Douglas-fir: state-of-the-art guidelines. Gen. Tech. Rep. INT-198. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 19 p. [1050]
101. Gullion, Gordon W. 1964. Contributions toward a flora of Nevada. No. 49: Wildlife uses of Nevada plants. CR-24-64. Beltsville, MD: U.S. Department of Agriculture, Agricultural Research Service, National Arboretum Crops Research Division. 170 p. [6729]
102. Haeussler, S.; Coates, D.; Mather, J. 1990. Autecology of common plants in British Columbia: A literature review. Economic and Regional Development Agreement: FRDA Report 158. Victoria, BC: Forestry Canada, Pacific Forestry Centre; British Columbia Ministry of Forests, Research Branch. 272 p. [18033]
103. Hall, Frederick C. 1974. Key to some common forest-zone plants of northwestern Washington. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 34 p. [3235]
104. Halpern, Charles B.; Evans, Shelley A.; Nielson, Sarah. 1999. Soil seed banks in young, closed-canopy forests of the Olympic Peninsula, Washington: potential contributions to understory reinitiation. Canadian Journal of Botany. 77(7): 922-935. [33078]
105. Hamilton, Anthony; Archibald, W. Ralph. 1986. Grizzly bear habitat in the Kimsquit River Valley, coastal British Columbia: evaluation. In: Contreras, Glen P.; Evans, Keith E., compilers. Proceedings-grizzly bear habitat symposium; 1985 April 30 - May 2; Missoula, MT. Gen. Tech. Rep. INT-207. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 50-56. [10811]
106. Hamilton, E. H.; Peterson, L. D. 2006. Succession after slashburning in an Engelmann spruce-subalpine fir subzone variant: West Twin site. Tech. Rep. No. 028. Victoria, BC: British Columbia Ministry of Forests and range, Forest Science Program. 21 p. [64622]
107. Hamilton, E. 2006. Vegetation development and fire effects at the Walker Creek site: comparison of forest floor and mineral soil plots. Technical Report No. 026. Victoria, BC: British Columbia Ministry of Forests and Range, Forest Science Program. 28 p. [64621]
108. Hamilton, Evelyn H. 2006. Fire effects and post-burn vegetation development in the sub-boreal spruce zone: Mackenzie (Windy Point) Site, [Online]. Technical Report 033. Victoria, BC: Ministry of Forests and Range Forest Science Program (Producer). 19 p. Available: http://www.for.gov.bc.ca/hfd/pubs/Docs/Tr/Tr033.pdf [2008, October 1]. [64177]
109. Hamilton, Evelyn H. 2006. Vegetation response, fire effects, and tree growth after slashburning in the Engelmann spruce-subalpine fir zone: Goat River Site. Technical Report No. 037. Kamloops, BC: British Columbia Ministry of Forests and Range, Research Branch, Forest Science Program. 26 p. [66358]
110. Hamilton, Evelyn H.; Yearsley, H. Karen. 1988. Vegetation development after clearcutting and site preparation in the SBS zone. Economic and Regional Development Agreement: FRDA Report 018. Victoria, BC: Canadian Forestry Service, Pacific Forestry Centre; British Columbia Ministry of Forests and Lands. 66 p. [8760]
111. Hamilton, Evelyn; Peterson, Les. 2003. Response of vegetation to burning in a subalpine forest cutblock in central British Columbia: Otter Creek site. Research Report 23. Victoria, BC: British Columbia Ministry of Forests, Forest Science Program. 60 p. [46111]
112. Hanley, Thomas A.; Deal, Robert L.; Orlikowska, Ewa H. 2006. Relations between red alder composition and understory vegetation in young mixed forests of southeast Alaska. Canadian Journal of Forest Research. 36: 738-748. [63861]
113. Hann, Wendel; Havlina, Doug; Shlisky, Ayn; [and others]. 2005. Interagency fire regime condition class guidebook. Version 1.2, [Online]. In: Interagency fire regime condition class website. U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior; The Nature Conservancy; Systems for Environmental Management (Producer). Variously paginated [+ appendices]. Available: http://www.frcc.gov/docs/1.2.2.2/Complete_Guidebook_V1.2.pdf [2007, May 23]. [66734]
114. Harrington, Constance A. 2006. Biology and ecology of red alder. In: Deal, Robert L.; Harrington, Constance A., tech. eds. Red alder: a state of knowledge. Gen. Tech. Rep. PNW-GTR-669. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station: 21-43. [63767]
115. 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]
116. Heller, Christine A. 1953. Wild edible and poisonous plants of Alaska. College, AK: University of Alaska, Cooperative Agricultural Extension Service. 167 p. In cooperation with: U.S. Department of Agriculture. [37068]
117. Henderson, Jan A. 1978. Plant succession on the Alnus rubra/Rubus spectabilis habitat type in western Oregon. Northwest Science. 52(3): 156-167. [6393]
118. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992]
119. Hidayati, Siti N.; Baskin, Jerry M.; Baskin, Carol C. 2000. Morphological dormancy in seeds of two North American and one Eurasian species of Sambucus (Caprifoliaceae) with underdeveloped spatulate embryos. American Journal of Botany. 87(11): 1169-1678. [71072]
120. Hidayati, Siti Nur. 2000. A comparative study of seed dormancy in five genera of Caprifoliaceae. Lexington, KY: University of Kentucky. 247 p. Dissertation. [69084]
121. Hitchcock, C. Leo; Cronquist, Arthur; Ownbey, Marion. 1959. Vascular plants of the Pacific Northwest. Part 4: Ericaceae through Campanulaceae. Seattle, WA: University of Washington Press. 510 p. [1170]
122. Hosley, N. W.; Ziebarth, R. K. 1935. Some winter relations of the white-tailed deer to the forests in north central Massachusetts. Ecology. 16(4): 535-553. [64485]
123. Hudson, Shelley; Carlson, Michael. 1998. Propagation of interior British Columbia native plants from seed. Victoria, BC: Ministry of Forests, Research Program. 30 p. [38690]
124. Hughes, Jeffrey W.; Cass, Wendy B. 1997. Pattern and process of a floodplain forest, Vermont, USA: predicted responses of vegetation to perturbation. Journal of Applied Ecology. 34(3): 594-612. [65684]
125. Hughes, Jeffrey W.; Fahey, Timothy J. 1991. Colonization dynamics of herbs and shrubs in disturbed northern hardwood forest. Journal of Ecology. 79: 605-616. [17724]
126. Hulten, Eric. 1968. Flora of Alaska and neighboring territories. Stanford, CA: Stanford University Press. 1008 p. [13403]
127. Hungerford, C. R. 1970. Response of Kaibab mule deer to management of summer range. Journal of Wildlife Management. 34(40): 852-862. [1219]
128. Hungerford, Roger D. 1984. Native shrubs: suitability for revegetating road cuts in northwestern Montana. Res. Pap. INT-331. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 13 p. [1220]
129. Hunt, Robert L. 1979. Removal of woody streambank vegetation to improve trout habitat. Tech. Bull. No. 115. Madison, WI: Department of Natural Resources. 37 p. [13744]
130. Jarvis, Robert L.; Passmore, Michael F. 1992. Ecology of band-tailed pigeons in Oregon. Biological Report 6. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service. 38 p. [64965]
131. Jenkins, Kurt J.; Starkey, Edward E. 1991. Food habits of Roosevelt elk. Rangelands. 13(6): 261-265. [17351]
132. Jicinska, Dagmar; Koncalova, Marie Nadezda. 1979. Flowering and fertilization process in European Sambucus and Quercus species. In: Bonner, F., ed. Proceedings: a synposium on flowreing and seed development in trees. Starkville, MS: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station: 103-111. [69006]
133. Johnson, Charles Grier, Jr. 1998. Vegetation response after wildfires in national forests of northeastern Oregon. R6-NR-ECOL-TP-06-98. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 128 p. [plus appendices]. [30061]
134. Kari, Priscilla Russell. 1987. Tanaina plantlore. Dena'ina K'et'una: An ethnobotany of the Dena'ina Indians of southcentral Alaska. 2nd ed. [Revised]. Anchorage, AK: U.S. Department of the Interior, National Park Service, Alaska Region. 205 p. [67343]
135. 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]
136. Kartesz, John Thomas. 1988. A flora of Nevada. Reno, NV: University of Nevada. 1729 p. [In 2 volumes]. Dissertation. [42426]
137. Kearney, Thomas H.; Peebles, Robert H.; Howell, John Thomas; McClintock, Elizabeth. 1960. Arizona flora. 2nd ed. Berkeley, CA: University of California Press. 1085 p. [6563]
138. Kearns, Carol Ann. 1990. The role of fly pollination in montane habitats. College Park, MD: University of Maryland. 208 p. Dissertation. [64729]
139. Kellman, M. C. 1970. The viable seed content of some forest soil in coastal British Columbia. Canadian Journal of Botany. 48: 1383-1385. [6469]
140. Kellman, Martin. 1974. Preliminary seed budgets for two plant communities in coastal British Columbia. Journal of Biogeography. 1: 123-133. [71054]
141. Kelly, George W. 1970. A guide to the woody plants of Colorado. Boulder, CO: Pruett Publishing Co. 180 p. [6379]
142. 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]
143. 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]
144. 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]
145. Kollmann, Johannes. 1995. Regeneration window for fleshy-fruited plants during scrub development on abandoned grassland. Ecoscience. 2(3): 213-222. [69004]
146. Kotar, John; Kovach, Joseph A.; Locey, Craig T. 1988. Field guide to forest habitat types of northern Wisconsin. Madison, WI: University of Wisconsin, Department of Forestry; Wisconsin Department of Natural Resources. 217 p. [11510]
147. 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]
148. Kraft, Lidia Szabo; Crow, Thomas R.; Buckley, David S.; Nauertz, Elizabeth A.; Zasada, John C. 2004. Effects of harvesting and deer browsing on attributes of understory plants in northern hardwood forests, Upper Michigan, USA. Forest Ecology and Management. 199(2-3): 219-230. [68986]
149. Kramer, Neal B. 1984. Mature forest seed banks on three habitat types in central Idaho. Moscow, ID: University of Idaho. 106 p. Thesis. [1375]
150. Kudish, Michael. 1992. Adirondack upland flora: an ecological perspective. Saranac, NY: The Chauncy Press. 320 p. [19376]
151. 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]
152. Lafferty, R. R. 1972. Regeneration and plant succession as related to fire intensity on clear-cut logged areas in coastal cedar-hemlock type: an interim report. Internal Report BC-33. Victoria, BC: Department of the Environment, Canadian Forestry Service, Pacific Forest Research Centre. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Lab, Missoula, MT. 129 p. [9985]
153. LANDFIRE Rapid Assessment. 2005. Potential Natural Vegetation Group (PNVG) R1PICOcw--Sierra Nevada lodgepole pine - cold wet upper montane, [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: http://www.landfire.gov/zip/CA/R1PICOcw_Aug08.pdf [2008, October 24]. [71583]
154. LANDFIRE Rapid Assessment. 2005. Potential Natural Vegetation Group (PNVG) R6JAPlop--Great Lakes pine barrens, [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: http://www.landfire.gov/zip/GL/R6JAPlop_Aug08.pdf [2008, October 24]. [71584]
155. LANDFIRE Rapid Assessment. 2005. Potential Natural Vegetation Group (PNVG) R7NESF--northeast spruce-fir forest, [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: http://www.landfire.gov/zip/NE/R7NESF_Aug08.pdf [2008, October 24]. [71581]
156. LANDFIRE Rapid Assessment. 2005. Potential Natural Vegetation Group (PNVG) R7NHNE--northern hardwoods northeast, [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: http://www.landfire.gov/zip/NE/R7NHNE_Aug08.pdf [2008, October 24]. [71580]
157. LANDFIRE Rapid Assessment. 2005. Potential Natural Vegetation Group (PNVG) R7NHSP--northern hardwoods-spruce, [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: http://www.landfire.gov/zip/Ne/R7NHSP_Aug08.pdf [2008, October 24]. [71582]
158. LANDFIRE Rapid Assessment. 2005. Potential Natural Vegetation Group (PNVG) R8PIECap--Appalachian shortleaf pine, [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: http://www.landfire.gov/zip/SA/R8PIECap_Aug08.pdf [2008, October 24]. [71579]
159. LANDFIRE Rapid Assessment. 2005. Reference condition modeling manual (Version 2.1), [Online]. In: LANDFIRE. Cooperative Agreement 04-CA-11132543-189. Boulder, CO: The Nature Conservancy; U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior (Producers). 72 p. Available: http://www.landfire.gov/downloadfile.php?file=RA_Modeling_Manual_v2_1.pdf [2007, May 24]. [66741]
160. 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: http://www.landfire.gov/models_EW.php [2008, April 18] [66533]
161. Larsen, J. A. 1923. Association of trees, shrubs, and other vegetation in the northern Idaho forests. Ecology. 4(1): 63-67. [60168]
162. Love, Askell; Love, Doris. 1954. Vegetation of a prairie marsh. Bulletin of the Torrey Botanical Club. 81(1): 16-34. [18103]
163. Lyon, L. Jack. 1966. Initial vegetal development following prescribed burning of Douglas-fir in south-central Idaho. Res. Pap. INT-29. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 17 p. [1494]
164. Lyon, L. Jack. 1984. The Sleeping Child Burn--21 years of postfire change. Res. Pap. INT-330. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 17 p. [6328]
165. 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]
166. Mabry, Cathy; Korsgren, Tobe. 1998. A permanent plot study of vegetation and vegetation-site factors fifty-three years following disturbance in central New England, U.S.A. Ecoscience. 5(2): 232-240. [45981]
167. MacLean, David A.; Wein, Ross W. 1977. Changes in understory vegetation with increasing stand age in New Brunswick forests: species composition, cover, biomass, and nutrients. Canadian Journal of Botany. 55: 2818-2831. [10106]
168. Manning W. J.; Godzik, B. 2004. Bioindicator plants for ambient ozone in Central and Eastern Europe. Environmental Pollution. 130(1): 33-39. [68987]
169. Marcum, C. Les. 1975. Summer-fall habitat selection and use by a western Montana elk herd. Missoula, MT: University of Montana. 188 p. Dissertation. [51342]
170. Martin, Alexander C.; Zim, Herbert S.; Nelson, Arnold L. 1951. American wildlife and plants. New York: McGraw-Hill Book Company, Inc. 500 p. [4021]
171. Martin, William H. 1990. The role and history of fire in the Daniel Boone National Forest. Final Report. Winchester, KY: U.S. Department of Agriculture, Forest Service, Daniel Boone National Forest. 131 p. [43630]
172. McCutcheon, A. R.; Roberts, T. E.; Gibbons, E.; Ellis, S. M.; Babiuk, L. A.; Hancock, R. E. W.; Towers, G. H. N. 1995. Antiviral screening of British Columbian medicinal plants. Journal of Ethnopharmacology. 49(2): 101-110. [68990]
173. McGee, Gregory G. 2001. Stand-level effects on the role of decaying logs as vascular plant habitat in Adirondack northern hardwood forests. Journal of the Torrey Botanical Society. 128(4): 370-380. [66831]
174. Meehan, William R. 1974. The forest ecosystem of southeast Alaska: 4. Wildlife habitats. Gen. Tech. Rep. PNW-16. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 32 p. [13479]
175. Metzger, Fred; Schultz, Jan. 1984. Understory response to 50 years of management of a northern hardwood forest in Upper Michigan. The American Midland Naturalist. 112(2): 209-223. [64467]
176. Mitchell, W. W. 1968. On the ecology of Sitka alder in the subalpine zone of south-central Alaska. In: Trappe, J. M.; Franklin, J. F.; Tarrant, R. F.; Hansen, G. M., eds. Biology of alder: Proceedings of a symposium; 1967 April 14-15; Pullman, WA. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station: 45-56. [17733]
177. Mladenoff, David J. 1990. The relationship of the soil seed bank and understory vegetation in old-growth northern hardwood-hemlock treefall gaps. Canadian Journal of Botany. 68: 2714-2721. [13477]
178. Mohlenbrock, Robert H. 1986. [Revised edition]. Guide to the vascular flora of Illinois. Carbondale, IL: Southern Illinois University Press. 507 p. [17383]
179. Monsen, Stephen B.; Stevens, Richard; Shaw, Nancy L. 2004. Shrubs of other families. 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: 598-698. [52846]
180. 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]
181. Morris, William G. 1970. Effects of slash burning in overmature stands of the Douglas-fir region. Forest Science. 16(3): 258-270. [4810]
182. 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]
183. Mueggler, Walter F. 1965. Ecology of seral shrub communities in the cedar-hemlock zone of northern Idaho. Ecological Monographs. 35: 165-185. [4016]
184. Mueggler, Walter F.; Campbell, Robert B., Jr. 1986. Aspen community types of Utah. Res. Pap. INT-362. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 69 p. [1714]
185. Munda, P.; Pater, M. 2001. Commercial sources of conservation plant materials, [Online]. Tucson, AZ: U.S. Department of Agriculture, Natural Resources Conservation Service, Tucson Plant Materials Center (Producer). Available: http://plant-materials.nrcs.usda.gov/pubs/azpmsarseedlist0501.pdf [2003, August 25]. [44989]
186. Munz, Philip A.; Keck, David D. 1973. A California flora and supplement. Berkeley, CA: University of California Press. 1905 p. [6155]
187. Neiland, Bonita J. 1958. Forest and adjacent burn in the Tillamook Burn area of northwestern Oregon. Ecology. 39(4): 660-671. [8879]
188. Netzel, M.; Strass, G.; Herbst, M.; Dietrich, H.; Bitsch, R.; Bitsch, I.; Frank, T. 2005. The excretion and biological antioxidant activity of elderberry antioxidants in healthy humans. Food Research International. 38(8-9): 905-910. [68985]
189. Norton, H. H.; Hunn, E. S.; Martinsen, C. S.; Keely, P. B. 1984. Vegetable food products of the foraging economies of the Pacific Northwest. Ecology of Food and Nutrition. 14(3): 219-228. [10327]
190. Pabst, Robert J.; Spies, Thomas A. 1998. Distribution of herbs and shrubs in relation to landform and canopy cover in riparian forests of coastal Oregon. Canadian Journal of Botany. 76: 298-315. [28627]
191. Park, Barry C. 1942. The yield and persistence of wildlife food plants. Journal of Wildlife Management. 6(2): 118-121. [7446]
192. 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]
193. Plummer, A. Perry. 1977. Revegetation of disturbed Intermountain area sites. In: Thames, J. C., ed. Reclamation and use of disturbed lands of the Southwest. Tucson, AZ: University of Arizona Press: 302-337. [27411]
194. Radford, Albert E.; Ahles, Harry E.; Bell, C. Ritchie. 1968. Manual of the vascular flora of the Carolinas. Chapel Hill, NC: The University of North Carolina Press. 1183 p. [7606]
195. Ralphs, M. H.; Pfister, J. A. 1992. Cattle diets in tall forb communities on mountain ranges. Journal of Range Management. 45(6): 534-537. [20189]
196. Ramovs, B. V.; Roberts, M. R. 2005. Response of plant functional groups within plantations and naturally regenerated forests in southern New Brunswick, Canada. Canadian Journal of Forest Research. 35(6): 1261-1276. [61144]
197. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]
198. Reagan, Albert B. 1934. Plants used by the Hoh and Quileute Indians. Transactions of the Kansas Academy of Science. 37: 55-70. [66487]
199. Ritter, C. M.; McKee, G. W. 1964. The elderberry, history, classification and culture. Pa. Agr. Exp. Sta. Bull. 709 22 pgs. [1221]
200. Roberts, Catherine Anne. 1975. Initial plant succession after brown and burn site preparation on an alder-dominated brushfield in the Oregon Coast Range. Corvallis, OR: Oregon State University. 90 p. Thesis. [9786]
201. Roberts, Warren G.; Howe, J. Greg; Major, Jack. 1980. A survey of riparian forest flora and fauna in California. In: Sands, Anne, ed. Riparian forests in California: Their ecology and conservation: Symposium proceedings; 1977 May 14; Davis, CA. Institute of Ecology Publication No. 15. Davis, CA: University of California, Division of Agricultural Sciences: 3-19. [5271]
202. Rogers, Robert S. 1980. Hemlock stands from Wisconsin to Nova Scotia: transitions in understory composition along a floristic gradient. Ecology. Vol. 61, No. 1: 178-193. [62813]
203. Rogers, Robert S. 1981. Mature mesophytic hardwood forest: community transitions, by layer, from east-central Minnesota to southeastern Michigan. Ecology. 62(6): 1634-1647. [22184]
204. Roland, A. E.; Smith, E. C. 1969. The flora of Nova Scotia. Halifax, NS: Nova Scotia Museum. 746 p. [13158]
205. Sampson, Arthur W.; Jespersen, Beryl S. 1963. California range brushlands and browse plants. Berkeley, CA: University of California, Division of Agricultural Sciences, California Agricultural Experiment Station, Extension Service. 162 p. [3240]
206. Saunders, Paul R.; Smathers, Garrett A.; Ramseur, George S. 1983. Secondary succession of a spruce-fir burn in the Plott Balsam Mountains, North Carolina. Castanea. 48(1): 41-47. [8658]
207. Schreiner, Edward G.; Krueger, Kirsten A.; Happe, Patricia J.; Houston, Douglas B. 1996. Understory patch dynamics and ungulate herbivory in old-growth forests of Olympic National Park, Washington. Canadian Journal of Forest Research. 26: 255-265. [26590]
208. 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]
209. Soper, James H.; Heimburger, Margaret L. 1982. Shrubs of Ontario. Life Sciences Miscellaneous Publications. Toronto, ON: Royal Ontario Museum. 495 p. [12907]
210. Stewart, R. E. 1978. Origin and development of vegetation after spraying and burning in a coastal Oregon clearcut. Res. Note PNW-317. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 11 p. [6541]
211. Stewart, Robert E. 1944. Food habits of blue grouse. The Condor. 46(3): 112-120. [55518]
212. Stickney, Peter F. 1980. Data base for post-fire succession, first 6 to 9 years, in Montana larch-fir forests. Gen. Tech. Rep. INT-62. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 133 p. [6583]
213. 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]
214. Stickney, Peter F. 1989. Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. FEIS workshop: Postfire regeneration. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 10 p. [20090]
215. 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]
216. 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]
217. Stiles, Edmund W. 1980. Patterns of fruit presentation and seed dispersal in bird-disseminated woody plants in the eastern deciduous forest. The American Naturalist. 116(5): 670-688. [6508]
218. Strausbaugh, P. D.; Core, Earl L. 1977. Flora of West Virginia. 2nd ed. Morgantown, WV: Seneca Books, Inc. 1079 p. [23213]
219. Stutchbury, Bridget J. M.; Capuano, Bianca; Fraser, Gail S. 2005. Avian frugivory on a gap-specialist, the red elderberry (Sambucus racemosa). Wilson Bulletin. 117(4): 336-340. [68984]
220. Suring, Lowell H.; Goldstein, Michael I.; Howell, Susan; Nations, Christopher S. 2006. Effects of spruce beetle infestations on berry productivity on the Kenai Peninsula, Alaska. Forest Ecology and Management. 227(3): 247-256. [62653]
221. Swan, Frederick Robbins, Jr. 1966. The effects of fire on plant communities of south-central New York State. Ithaca, NY: Cornell University. 169 p. Dissertation. [37434]
222. Swetnam, Thomas W.; Baisan, Christopher H. 1996. Historical fire regime patterns in the southwestern United States since AD 1700. In: Allen, Craig D., ed. Fire effects in Southwestern forests: Proceedings, 2nd La Mesa fire symposium; 1994 March 29-31; Los Alamos, NM. RM-GTR-286. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 11-32. [27281]
223. Talley, Theresa Sinicrope; Fleishman, Erica; Holyoak, Marcel; Murphy, Dennis D.; Ballard, Adam. 2007. Rethinking a rare-species conservation strategy in an urban landscape: the case of the valley elderberry longhorn beetle. Biological Conservation. 135(1): 21-32. [68997]
224. Taylor, Dale L. 1969. Biotic succession of lodgepole pine forests of fire origin in Yellowstone National Park. Laramie, WY: University of Wyoming. 320 p. Thesis. [9481]
225. Taylor, R. F. 1932. The successional trend and its relation to second-growth forests in southeastern Alaska. Ecology. 13(4): 381-391. [10007]
226. Thilenius, John F. 1990. Woody plant succession on earthquake-uplifted coastal wetlands of the Copper River Delta, Alaska. Forest Ecology and Management. 33/34: 439-462. [11803]
227. 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]
228. Traveset, Anna; Willson, Mary F. 1997. Effect of birds and bears on seed germination of fleshy-fruited plants in temperate rainforests of southeast Alaska. Oikos. 80(1): 89-95. [67728]
229. Tremblay, Nicolas O.; Larocque, Guy R. 2001. Seasonal dynamics of understory vegetation in four eastern Canadian forest types. International Journal of Plant Science. 162(2): 271-286. [47261]
230. Turner, Nancy Chapman; Bell, Marcus A. M. 1973. The ethnobotany of the southern Kwakiutl Indians of British Columbia. Economic Botany. 27: 257-310. [21015]
231. 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]
232. U.S. Department of Agriculture, Forest Service. 1937. Range plant handbook. Washington, DC. 532 p. [2387]
233. U.S. Department of Agriculture, Natural Resources Conservation Service. 2008. PLANTS Database, [Online]. Available: http://plants.usda.gov/. [34262]
234. U.S. Department of the Interior, Fish and Wildlife Service, Division of Endangered Species. 2008. Threatened and endangered animals and plants, [Online]. Available: http://www.fws.gov/endangered/wildlife.html [2008, September 23]. [62042]
235. Usui, Masayuki; Kakuda, Yukio; Kevan, Peter G. 1994. Composition and energy values of wild fruits from the boreal forest of northern Ontario. Canadian Journal of Plant Science. 74(3): 581-587. [24583]
236. Viereck, Leslie A.; Little, Elbert L., Jr. 1972. Alaska trees and shrubs. Agric. Handb. 410. Washington, DC: U.S. Department of Agriculture, Forest Service. 265 p. [6884]
237. Vogl, Richard J.; Ryder, Calvin. 1969. Effects of slash burning on conifer reproduction in Montana's Mission Range. Northwest Science. 43(3): 135-147. [8546]
238. Voss, Edward G. 1996. Michigan flora. Part III: Dicots (Pyrolaceae--Compositae). Bulletin 61: Cranbrook Institute of Science; University of Michigan Herbarium. Ann Arbor, MI: The Regents of the University of Michigan. 622 p. [30401]
239. Weber, William A.; Wittmann, Ronald C. 1996. Colorado flora: eastern slope. 2nd ed. Niwot, CO: University Press of Colorado. 524 p. [27572]
240. 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]
241. 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]
242. Winterhalder, Keith. 1990. The trigger-factor approach to the initiation of natural regeneration of plant communities on industrially-damaged lands at Sudbury, Ontario. In: Hughes, H. Glenn; Bonnicksen, Thomas M., eds. Restoration '89: the new management challenge: Proceedings, 1st annual meeting of the Society for Ecological Restoration; 1989 January 16-20; Oakland, CA. Madison, WI: The University of Wisconsin Arboretum, Society for Ecological Restoration: 215-226. [14697]
243. 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]
244. Wofford, B. Eugene. 1989. Guide to the vascular plants of the Blue Ridge. Athens, GA: The University of Georgia Press. 384 p. [12908]
245. Worley, D. M.; Nixon, C. M. 1974. Elders. In: Gill, J. D.; Healy, W. M., compilers. Shrubs and vines for northeastern wildlife. Gen. Tech. Rep. NE-9. Upper Darby, PA: U. S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 48-51. [6707]
246. Wright, Henry A. 1972. Shrub response to fire. 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: 204-217. [2611]
247. Yorks, Thad E.; Leopold, Donald J.; Raynal, Dudley J. 2000. Vascular plant propagule banks of six eastern hemlock stands in the Catskill Mountains of New York. Journal of the Torrey Botanical Society. 127(1): 87-93. [37018]
248. Zager, Peter Edward. 1980. The influence of logging and wildfire on grizzly bear habitat in northwestern Montana. Missoula, MT: University of Montana. 131 p. Dissertation. [5032]
249. Zamora, Benjamin Abel. 1975. Secondary succession on broadcast-burned clearcuts of the Abies grandis-Pachistima myrsinites habitat type in northcentral Idaho. Pullman, WA: Washington State University. 127 p. Dissertation. [5154]
250. Zobel, Donald B. 2002. Ecosystem use by indigenous people in an Oregon coastal landscape. Northwest Science. 76(4): 304-314. [64344]

FEIS Home Page