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Sambucus racemosa

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

INTRODUCTORY

AUTHORSHIP AND CITATION FEIS ABBREVIATION NRCS PLANT CODE COMMON NAMES TAXONOMY SYNONYMS LIFE FORM FEDERAL LEGAL STATUS OTHER STATUS
	Red elderberry along Turnagain Arm near Bird, Alaska. Photo by Mary Hopson.

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.

DISTRIBUTION AND OCCURRENCE

• GENERAL DISTRIBUTION
• HABITAT TYPES AND PLANT COMMUNITIES

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.

• Rocky Mountain elderberry is an indicator species of grand fir (Abies grandis) mosaic habitat types of northern Idaho. It is among the few woody plants that grow in western bracken fern (Pteridium aquilinum) glades within grand fir forests [82].

• Rocky Mountain elderberry riparian vegetation type [18]

• quaking aspen (Populus tremuloides)/red elderberry community type of mid- to high elevations (8,000-9,000 feet (2,440-2,740 m)) [183]

• sweet birch-eastern hemlock-American beech (Betula lenta-Tsuga canadensis-Fagus grandifolia)/red elderberry/roughleaf ricegrass (Oryzopsis asperifolia) northern mesic forest type [58]

• red alder-salmonberry (Rubus spectabilis)-red elderberry floodplain association of the Kimsquit River valley; a seral stage of the black cottonwood-Sitka spruce (Populus balsamifera subsp. trichocarpa-Picea sitchensis)/salmonberry association [15]

• sugar maple (Acer saccharum)-American beech/striped maple-alternate-leaf dogwood (A. pensylvanicum-Cornus alternifolia)/red elderberry plant association of the St Lawrence Valley [15]

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

GENERAL BOTANICAL CHARACTERISTICS REGENERATION PROCESSES SITE CHARACTERISTICS SUCCESSIONAL STATUS SEASONAL DEVELOPMENT
	National Park Service photo by Richard Thomas.

• Botanical description
• Raunkiaer life form

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].

• Vegetative regeneration
• Pollination and breeding system
• Flower, fruit, and seed production
• Seed dispersal
• Seed banking
• Germination
• Seedling establishment and plant growth

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]:

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:

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.

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].

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].

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].

FIRE EFFECTS AND MANAGEMENT

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

• Immediate fire effect on plant
• Postfire regeneration strategy
• Fire adaptations and plant response to fire

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.

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.

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].

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].

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].

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]:

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.

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
• Fire regimes

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. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".

MANAGEMENT CONSIDERATIONS

	IMPORTANCE TO WILDLIFE AND LIVESTOCK VALUE FOR REHABILITATION OF DISTURBED SITES OTHER USES OTHER MANAGEMENT CONSIDERATIONS
Photo used with permission of Creative Commons.

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].

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].

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].

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

APPENDIX: FIRE REGIME TABLE

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