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Rubus spectabilis, salmonberry
Figure 1—Salmonberry flower.
Photo by Barry Breckling ©2015.
Figure 2—Salmonberry patch in flower.
Photo by Jean Pawek ©2011.

Citation:
Zouhar, Kris. 2019. Rubus spectabilis, salmonberry. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Missoula Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/plants/plants/shrub/rubspe/all.html [].


Table of Contents


SUMMARY


This review summarizes information that was available in the scientific literature as of 2019 on the biology, ecology, and effects of fire on salmonberry in North America.

Available literature on salmonberry is dominated by botanical descriptions, reports on habitat preferences and associated species in floras and plant community classifications, silviculture articles about its competition with conifer seedlings, and reports on control by herbicides. Several publications address salmonberry regeneration strategies and successional roles. Few publications were available that specifically addressed salmonberry response to fire, with the exception of those describing its response to slashburning.

Salmonberry reproduces vegetatively via buds located on rhizomes, root crowns, stumps, and possibly from aerial stems. It also reproduces from animal- and gravity-dispersed seed, and forms a persistent seed bank. On undisturbed sites individual clones and populations increase in cover primarily by aerial stems sprouting from rhizomes. Recruitment of new plants through seedling establishment is relatively rare, but it is the primary means by which salmonberry establishes on new sites. Seedlings may also establish from the soil seed bank after severe soil disturbance. Although disturbance is not necessary for maintaining dense cover of salmonberry, it sprouts prolifically after aerial stems are killed and after overstory trees die or are removed.

Salmonberry stands annually produce new aerial stems and new rhizomes, which maintain dense cover, given adequate light and moisture. Salmonberry population growth and clone size are influenced by density of overstory trees, stand type, moisture availability, disturbance severity, and salmonberry basal area. The largest and most dominant populations of salmonberry occur on sites with a history of major disturbance, such as after logging, wildfire, abandonment of agricultural land, and along waterways and roadsides.

Salmonberry has relatively high shade tolerance compared to other Rubus species, but does not grow well in deep shade. It is most abundant in early secondary succession, and can grow and spread rapidly after canopy removal. It generally increases in cover after overstory removal; increases are mostly from sprouting, depending on severity of understory and soil disturbance. It is tolerant of frequent flooding that occurs in some riparian communities, and it may form persistent shrub communities in these areas.

Salmonberry reproductive traits (i.e., buried seed, buried rhizomes, and rapid sprouting following top-kill) make it moderately to highly resistant to fire. Rhizomes and root crowns generally survive even when aerial stems are killed. Salmonberry can recover rapidly after fire through both sprouting and seedling establishment. Actual postfire plant mortality is presumably low. Seed in the soil seed bank is probably unharmed by fire, while that in the litter and duff may be killed by severe fire that consumes surface organic layers. Severe fire, particularly on dry sites, may reduce salmonberry sprouting potential and canopy cover in the short term (~10 years).


INTRODUCTORY


TAXONOMY
The scientific name of salmonberry is Rubus spectabilis Pursh (Rosaceae) [16,73,139,145,157].

The PLANTS Database (2019) recognizes two varieties, both with the common name salmonberry:

Salmonberry hybridizes with grayleaf raspberry and arctic raspberry in Alaska [238,330].

Common names are used throughout this review. For scientific names and links to other FEIS Species Reviews, see table A2 for plants and table A4 for animals.

SYNONYMS
None

LIFE FORM
Shrub-liana

DISTRIBUTION AND OCCURRENCE

SPECIES: Rubus spectabilis
GENERAL DISTRIBUTION
Coarse distribution of salmonberry. Map courtesy of the PLANTS Database (2019, 8 May) [326].

Salmonberry is native to the Pacific coast states and Idaho. It grows mostly west of the Cascade Range in Washington and Oregon southward to northwestern California [102,267] and along the Pacific Coast northward through coastal British Columbia to southern Alaska and the Aleutian Islands [18,139,274,275]. Its frequency generally declines from west to east [63]. It is uncommon east of the Cascade crest [102], where its distribution often follows watercourses [102] such as the banks of the Columbia River [145]. Disjunct populations occur in Idaho, although they are rare [18,148]. NatureServe (2019) reports its occurrence in West Virginia [227].

Rubus spectabilis var. spectabilis occurs in Mendocino, Humboldt, and Del Norte counties in California, northward to Alaska, and eastward to Idaho [326]. Rubus spectabilis var. franciscanus occurs only in Oregon and along the Pacific Coast in California, from the Santa Cruz Mountains north to Sonoma County [18,326].

Salmonberry's distribution is limited by cold temperatures and short growing seasons, and it is restricted primarily to mild maritime climates [238], decreasing in abundance inland (e.g., [316]) and where the climate has more continental influence [258].

States and provinces [227,326]:
United States: AK, CA, ID, OR, WA
Canada: BC

SITE CHARACTERISTICS
Salmonberry is most common on moist to wet, water-receiving sites in forested or wooded areas, especially in openings, and along edges and streambanks [16,139,145,238,251,275,324], although it also occurs on relatively dry hillsides and disturbed areas [18]. It is an indicator of warm, wet sites (e.g., [3,201]), is common in riparian areas (e.g., [315]), and is a facultative wetland plant (e.g., [176,201]). It typically occurs in forest openings, along waterways, on river terraces, gravel bars, avalanche chutes, or in seeps and swamps [42,75,102,139,251,330]. Its growth may be reduced in excessively wet areas such as swamps, where it may be confined to hummocks or logs [18]. Salmonberry can be abundant in disturbed areas such as roadsides, fencerows, fallow fields, and logged or burned areas [18,43,324].

Salmonberry typically grows at low to middle elevations [139]. Its frequency of occurrence decreases with increasing elevation [258], and it is generally most abundant below about 2,600 feet (800 m) [238]. However, it occurs up to lower alpine elevations in Alaska [145], and to subalpine elevations in the Pacific Northwest [251]. It occurs from sea level to >4,000 feet (1,200 m) in the Cascade and Coast ranges of Washington and Oregon [324]. In western Washington, it is particularly abundant under forest canopies at lower elevations but is largely restricted to stream and lake margins at higher elevations [18]. It is most abundant below 3,000 feet (900 m) in Oregon [316] and below about 1,600 feet (500 m) in California [16].

Although it may tolerate a range of soil types [18], salmonberry is most abundant and attains maximum size in rich, moist soils of alluvial bottomlands [324]. It is a nitrophytic species and an indicator of nutrient-rich [169,171,238,258] and very moist to wet soils in forests of British Columbia [258]. In western Oregon and Washington, salmonberry is an indicator of fertile soils that are saturated for much of the year [102,189]. In the Coast Ranges of western Oregon, the Douglas-fir/salmonberry–western swordfern community had the greatest nutrient concentrations compared to Douglas-fir communities with other understory types due to the presence of red alder [346]—a nitrogen-fixing species and common associate of salmonberry (see Plant Communities and Succession). In British Columbia, salmonberry was an indicator of fertile sites where black cottonwood grew best [284].

For additional information on soil characteristics on sites occupied or dominated by salmonberry, see table A3 for a list of vegetation classifications, most of which include information on soils and other site characteristics. Also see table 1 for information on studies that examined relationships between site characteristics and vegetation, including salmonberry.

Table 1—Publications with information on salmonberry site characteristics in specific locations.
Location Information Citation
Alaska
Chugach National Forest, Kenai Peninsula Evaluated cover of berry-producing species relative to landscape position and environmental conditions, including crown closure and spruce beetle infestation. Suring et al. (2008) [301]
British Columbia
University of British Columbia Research Forest and Vancouver Island Characterized the relationships among humus (soil organic matter) forms, forest floor nutrient properties, and understory vegetation at 151 cool, mesothermal forest sites Klinka et al. (1990) [171]
southwestern British Columbia Examined potential for estimating site index of Douglas-fir from ecological variables Green et al. (1989) [98]
eastern Vancouver Island and adjacent mainland Evaluated relationships between site index, plant communities, and site characteristics in coastal Douglas-fir ecosystems Klinka et al. (1989) [165]
southwestern British Columbia Characterized the most productive ecosystems for Douglas-fir growth Klinka et al. (1981) [166]
southwestern British Columbia Characterized the most productive ecosystems Engelmann spruce growth Klinka et al. (1982) [167]
Oregon
Four major watersheds in western Oregon Analyses of woody riparian plant distribution and diversity across a climate-productivity gradient at multiple scales Sarr and Hibbs (2007) [271]
Southern Coast Ranges Characterized gradients of plant species composition and groups of species associated with specific geomorphic areas in 63 zero-order basins Sheridan and Spies (2005) [281]
Cummins Creek watershed, Coast Ranges

Associated salmonberry with moist areas and riparian areas (by regression) in Sitka spruce-western hemlock and Douglas-fir forests.

Wimberly and Spies (2001) [342]
Alsea River watershed Characterized the distribution of shrub and herb species relative to landform and forest canopy attributes at 94 sites along 42 streams

Pabst and Spies (1998) [243]

Siuslaw National Forest

Provided height growth and site index curves for Douglas-fir

Means and Sabin (1989) [209]
California
Humboldt Redwoods State Park Vegetation occurrence in relation to gradients of moisture, nutrients, light, and temperature Waring and Major (1964) [337]

PLANT COMMUNITIES
Salmonberry occurs in deciduous hardwood, mixed hardwood-coniferous evergreen, and coniferous forests and woodlands. It tends to be more frequent and develop greater canopy cover under red alder stands than in conifer or mixed hardwood-conifer stands (e.g., [18,83,307]). Salmonberry is a common to dominant understory species in coniferous forests dominated by Sitka spruce [125], redwood [194], western redcedar, western hemlock, Douglas-fir [219], Port-Orford-cedar, grand fir [352], and Pacific silver fir [75,76,188,351]. Salmonberry often forms dense patches within the understories of Douglas-fir and western hemlock forests [308]. It is especially common under red alder on both upland and riparian sites (e.g., [80,104,116]). In the southern portion of its range, salmonberry is considered an especially good indicator of potential red alder habitat [117]. It is a common component of northern coastal scrub communities [292] dominated by baccharis, thimbleberry, and trailing blackberry in northern California [128]. Salmonberry also grows in mixed-evergreen and hardwood forests [18,225,338] and in riparian forests dominated by black cottonwood [60], Sitka alder, and other hardwoods [77]. In Alaska, salmonberry commonly occurs in the understory of Sitka alder thickets [175].

Salmonberry has been identified as a dominant species under Sitka spruce, western hemlock, western redcedar, mountain hemlock, red alder, Sitka alder, California hazelnut, vine maple, and Oregon ash; and a codominant with these shrubs: salal, stink currant, thimbleberry, California blackberry, California huckleberry, devilsclub, common snowberry, northern black currant, and Alaska blueberry. Common understory associates include the forbs seacoast angelica, threeleaf foamflower, coastal miterwort, American skunkcabbage, Pacific golden saxifrage, and Pacific waterleaf; the graminoid bluejoint; and the ferns lady-fern and western swordfern (e.g., [21,22,64,66,82,130,172,176,201]).

For a list of species that commonly occur with salmonberry and links to available FEIS Species Reviews, see table A2. For a list of vegetation classifications and related publications that include salmonberry as an important, indicator, or dominant species, see table A3.

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Rubus spectabilis
GENERAL BOTANICAL CHARACTERISTICS

Figure 3—Yellow salmonberry fruit, Thornton Creek watershed, Seattle, Washington. Photo by Ashley Pond 2006. Figure 4—Red salmonberry fruit. Photo by Vernon Smith ©2011.

Botanical Description
This description covers characteristics that may be relevant to fire ecology and is not meant for identification. Keys for identification are available (e.g., [16,139,145]).

Aboveground: Salmonberry is an erect [16,139,145,251], branching [225,251,330], perennial shrub that typically grows 7 to 13 feet (2-4 m) tall [16,225,251,316,330]. It may reach up to 15 feet (4.6 m) tall [102,189] and is rarely less than 1.6 feet (0.5 m) [139]. Vegetative characteristics of salmonberry stems are affected by light and other resource availability [309,347]. On hot, dry sites (e.g., open hillsides and clearcuts), salmonberry assumes a short, compact form, usually less than about 3 feet (1 m) tall [18]. Leaves are deciduous, alternate, and mostly with three leaflets [102,139,145,225,251,330]. Aerial stems are 0.1 to 0.6 inch (3-15 mm) in diameter, and often have scattered, weak spines [251,330]. As stems age, bark becomes shreddy [102,225,251,330]. Several sources report that salmonberry stems are biennial (e.g., [16,145,225,330]). However, this does not appear to be true. For example, stems at least 10 years old, and possibly 15 years old, have been observed in southern British Columbia and Oregon [99], and Zasada and Tappeiner (2008) indicate that salmonberry has relatively long-lived stems [347].

Salmonberry flowers are a showy, unusual magenta color, and they occur singly or in groups of two to four on short, leafy, slender lateral stalks [16,102,145,225,251,330]. Fruits are raspberry-like, round to ovoid [16,102,145,251], 0.6 to 0.8 inch (1.5-2 cm) in length [225], and made up of many small, glabrous drupelets [139,330] that each contain a single, hard-pitted seed [347]. Seeds are small, around 310 seeds/gram [345]. Fruits vary in color, from yellowish (fig. 3) to orange to deep red (fig. 4) [16,102,139,145]—a quality that has been studied for its effects on seed dispersal. Dominance and variation in fruit color may differ among sites and geographic areas. The orange form is generally more common in the southern part of its range (i.e., Oregon), and the red form in the northern part (i.e., southeastern Alaska), although plants with both fruit colors occur in both areas, and the red form passes through an orange stage as it matures (review by [347]).

Stand structure: Salmonberry clones are made up of a network of rhizomes connecting ramets that consist of a taproot and one to five aerial stems. Stands annually produce enough aerial stems and rhizomes to offset mortality and maintain persistent cover [238,305,309]. Rhizomes spread laterally and produce new aerial stems about 3 to 6 feet (1-2 m) from the parent plant [305]. The number of aerial stems decreases from small to large size classes [238,309,350].

Populations range from small and scattered patches to what are described as large, "dense thickets" [102,145,189,309,316,330], which may be "almost impenetrable" in lower alpine areas of Alaska [145]. Large populations typically occur in openings and along streams, and the largest populations of salmonberry occur on sites with a history of major disturbance such as logged or burned sites, abandoned agricultural land, and sites along waterways and roadsides [347]. Populations can range from several meters [99] to more than ~330 feet (100 m) in diameter [18]. Individual clones can cover areas up to about 377 feet2 (35 m2) [350], and density can exceed 8,000 aerial stems/acre (20,000 stems/ha) [305].

Salmonberry clone size (rhizome and aerial stem density and biomass) and population growth are influenced by density of overstory trees [197], stand type, disturbance severity [305,309], and salmonberry basal area [347]. In four common stand types in western Oregon—clearcuts, Douglas-fir, red alder, and riparian—salmonberry stem density was greatest in red alder stands and in clearcuts, and was lowest in riparian and Douglas-fir stands [305]. Salmonberry clones in red alder stands also had more total rhizome length (60 feet (18 m)) than those in Douglas-fir and riparian stands (16-20 feet (5-6 m)) [309]. In plots where all salmonberry stems were cut within 6 inches (15 cm) of the ground, stem production was generally two to three times greater than on uncut plots for 2 to 3 years after plot establishment (table 2). On uncut plots, mean annual salmonberry stem production was related to salmonberry basal area (P < 0.001), and salmonberry stem production, older stem density, and clonal biomass were inversely related to overstory basal area (P < 0.01) [305]. Most salmonberry aerial stem production and annual rhizome growth was in young (2-year-old) clearcuts, and total rhizome length and total above- and below-ground biomass were greater in older (13- to 18-year-old) clearcuts than in undisturbed red alder, Douglas-fir, and riparian stands. Ramet density was significantly greater in the young (11.0 ramets/m2) and older clearcuts (7.0 ramets/m2) than in the understory of alder (3.9 ramets/m2), Douglas-fir (2.0 ramets/m2), and riparian (2.3 ramets/m2) stands [309]. Salmonberry population structure, rhizome length, and biomass could be predicted from salmonberry stem number and basal area and basal area of overstory trees [238,309].

Table 2—Mean salmonberry aerial stem density at the beginning of study, and mean annual stem production (number of stems/m2) over 8 years in four stand types in the Oregon Coast Ranges [305].
Stand type (age) Mean stem density (range) Mean annual stem production
Uncut plots Cut plots*
Red alder (53-61 years) 5.3 (3-7) 5.1 ~7
Conifer (110-140+ years) 2.5 (2-4) 2.2 2.4
Riparian** (58-120+ years) 2.8 (1-5) ~2.5 3.4
Clearcut (4-6 years) 5.0 (3-8) 4.7 13.0
*All stems were cut to ground level in 1987, and new aerial stems were tallied each year from 1988 to 1996.
**Mixed-age stands of Douglas-fir, western hemlock, red alder, and bigleaf maple.

The following publications provide additional metrics describing clone morphology and stem production on different site types in the Oregon Coast Ranges: Tappeiner et al. (2001) [305], Maxwell et al. (1993) [198], Zasada et al. (1992) [350], Tappeiner et al. (1991) [309], and Maxwell (1990) [197].

Belowground: About 30% to 40% of total salmonberry biomass is below ground [309]. The extent of the rhizome network is related to overstory basal area and composition, age of the parent plant, and site characteristics [349]. Rhizome development is often extensive, especially in disturbed areas. Averages of 20 to 30 miles (32-48 km) per stand are common in logged areas, and may reach 42 miles (68 km) per stand in some clearcuts [349].

Belowground morphology differs among stand and site types in the Oregon Coast Ranges. The average rhizome length per clone was greatest in red alder stands (~60 feet (18.3 m)) (P < 0.05), compared to ~16 feet (5 m) in Douglas-fir stands, ~20 feet (6 m) in riparian stands, and ~26 feet (8 m) in 2-year-old clearcuts. In undisturbed stands (red alder, Douglas-fir, and riparian), rhizomes sprouted only from buds at the base of ramets, while in clearcuts rhizomes also sprouted from buds at several points along old rhizomes. Rhizome length and bud number can be estimated from regression equations using overstory basal area and density of salmonberry stems [309]. Salmonberry rhizomes supported many fine roots and occurred about 0.8 to 8 inches (2-20 cm) below the soil surface in study sites. Although the depth of a particular rhizome was generally consistent over its entire length, some varied in depth by as much as 8 inches (20 cm). On steep slopes (>50%) adjoining the study sites, rhizomes occurred at depths of about 3 to 6 feet (1-2 m). Rhizomes 2 to 3 inches (5-8 cm) in diameter were common on study sites, suggesting that rhizomes are long lived. Clones in undisturbed stands produced one to two new rhizomes per year, with annual extension ranging from 4 inches (10 cm) in riparian stands to 28 inches (70 cm) in red alder stands. Clones in recent clearcuts produced an average of seven new rhizomes per clone, with an annual extension of 6.2 feet (1.9 m), higher than in other stand types (P < 0.01) [309].

Raunkiaer [253] Life Form
Phanerophyte

SEASONAL DEVELOPMENT
Salmonberry buds may be active in late winter or very early spring, with bud burst and leaf flush occurring from March to April [18,238]. For example, researchers in western Washington observed the first salmonberry leaves by 21 March [18]. Leaves are generally fully expanded from May to late August [238].

Salmonberry flowering usually occurs from early to late spring but varies by geographic location and elevation (table 3). Flowering generally occurs between April and June in its southern range and between April and July in its northern range [238]. In many areas along the Pacific Coast, the time of flowering appears to coincide with the arrival of the migrating rufous hummingbird, which may be an important pollinator [250].

Table 3—Salmonberry flowering dates by location.
Location Dates
California February–June [16,41]
western Oregon and southwestern Washington March–June [102]
northwestern Oregon March–June [316]
Olympic National Forest March–June [189]
Alaska April–July [330]

Fruit ripening generally occurs 30 to 36 days after pollination, and it may last for several weeks or more within a population [347]. Salmonberry fruits generally ripen from June to July in its southern range and from July to August in its northern range and at high elevations [198,238]. Fruits ripen from May to July in Bella Coola, British Columbia [188], in July near Ketchikan, Alaska, and in August farther north [319,330]. Fruits may be available into October in southeastern Alaska [319]. Seed dispersal coincides with the time of fruit availability. In many areas, seed is dispersed from June through August [345].

Salmonberry leaves typically fall by late October or November [42,136,198,238]. In the western Cascade Range, 56% of salmonberry leaves dropped during the first 3 weeks after frost [42]. Salmonberry may become dormant in winter, or continue minimum shoot elongation throughout a mild coastal winter (e.g., mean temperature of ~43 °F (6 °C)) [18,238].

Rhizome growth begins in March, and the tips of terminal buds remain active and continue elongating until August [18]. Total nonstructural carbohydrates (TNC) of rhizome segments removed from intact plants reached a high of 13% of dry weight during winter (February), fell to 6% to 7% during spring shoot production and summer growth, and began rising again during leaf fall in October. Shoot production was more closely associated with TNC content than was root production. Low shoot production coincided with low levels of TNC, suggesting that salmonberry is more susceptible to physical disturbance from May through July, and possibly into October [348]. For more information on rhizome phenology see Zasada et al. (1994) [348].

REGENERATION PROCESSES

Salmonberry reproduces vegetatively via buds located on rhizomes, root crowns, and possibly from aerial stems. It also reproduces from animal- and gravity-dispersed seed. On undisturbed sites individual clones and populations increase in cover primarily by aerial stems sprouting from rhizomes. Recruitment of new genets through seedling establishment is relatively rare in existing salmonberry populations [309], but it is the primary means by which salmonberry establishes on new sites [238]. Although disturbance is not necessary for maintaining dense cover of salmonberry, it sprouts "vigorously" after aerial stems are killed [305,309] and after overstory trees die or are removed (e.g., [309]). Salmonberry populations on disturbed sites such as clearcuts are likely dominated by clonal sprouts rather than plants from seedlings [197].

Vegetative Regeneration: Salmonberry maintains stable populations through clonal spread and annual sprouting of new, relatively long-lived ramets [238,347]. Its stems arise from extensive bud banks, located primarily on branching rhizomes [102,189,198,238,251,309,330,347,348,350] that spread laterally beneath the soil surface, producing new aerial stems ~3 to 20 feet (1-6 m) away from the parent plant [99]. Annual production of aerial stems offsets aerial stem mortality and enables salmonberry to maintain persistent cover in disturbed and undisturbed stands [238,305,309], unless disturbance to salmonberry is severe enough to allow for succession of trees and other shrubs [238,309]. For example, salmonberry sprouting in riparian stands was likely reduced by establishment and spread of stink currant the first year after salmonberry stems were top-killed by cutting [305]. Vegetative regeneration is particularly important in perpetuating colonies in shaded understory habitats [18]. However, salmonberry may not sprout following top-kill in shaded habitats; it did not sprout after cutting in dense Douglas-fir stands [305].

Salmonberry stem production varies among stand types and is greatest in relatively open stands (e.g., red alder) and following overstory removal. Salmonberry stem density was maintained for at least 8 years, despite high annual mortality of small, new stems (85%-95%), in four common stand types in western Oregon: clearcuts, Douglas-fir, red alder, and riparian (mixed-age stands of Douglas-fir, western hemlock, red alder, and bigleaf maple). Few large diameter stems (>0.4 inch (11 mm)) died during the 8 years of this study. Based on length of rhizomes and bud density, researchers estimated that only 1% to 5% of the rhizome buds in a stand are needed to maintain annual stem production [305]. Within the first two growing seasons after removal of overstory trees, salmonberry populations were maintained by rapid growth of new rhizomes (1.0-2.5 m/m2 annually) and sprouting of aerial stems (25-50 stems/m2). Rhizome and aerial stem biomass were greater in 13- to 18-yr-old clearcuts, than in 2-year-old clearcuts, red alder, Douglas-fir, and riparian stands [309].

Salmonberry sprouts prolifically from buds on stumps, root crowns, and rhizomes following top-kill from fire, mechanical removal, and other types of disturbance [18,238,305,309,348,349]. In clearcuts, Douglas-fir, red alder, and riparian stands in the Oregon Coast Ranges, annual stem production was greater for 2 or 3 years on plots where all salmonberry stems were cut than on plots where stems were not cut. After the first few years, annual stem production was similar among all stand types. Mean annual stem production was somewhat greater on cut plots than uncut plots averaged over 8 years, especially in clearcut and red alder stands (table 2) [305].

Regeneration by layering (from buds on aerial stem tips) has also been reported [18,309].

Rhizomes: Rhizomes are the most extensive segment of the salmonberry bud bank. Sprouting rhizome buds are responsible for local, annual increases in aerial stem density [18,238,347,348] and salmonberry's potential to survive severe soil heating and soil disturbance [309,349]. Bud densities and sprouting potential seem to be greatest on young rhizomes; 1- to 2-year-old rhizomes may have bud densities as high as one to two buds per inch. These young rhizomes are actively growing and expanding the area occupied by salmonberry underground, and producing aerial stems when "conditions are right". Older, larger diameter rhizomes seem to have fewer living buds that are slower to sprout after top-kill [309,349].

Rhizome development is often extensive [309,349], but rhizome morphology varies among stand and site types (e.g., [305,309]). Salmonberry population structure, rhizome length, and total biomass are related to overstory density and composition and disturbance severity [197,305,309] and can be estimated from predisturbance overstory basal area, salmonberry stem number and basal area, and site characteristics [309].

Stumps: Buds present on the stump seem to have the greatest potential for immediate growth after top-kill, but they are also the most susceptible to mortality or removal from burning or cutting [349]. The number of buds remaining on stumps is largely determined by stump height. Stump sprouts may inhibit other less active buds located at or below the ground surface by establishing apical dominance [349].

Root crown: Buds on salmonberry root crowns occur at and below the soil surface, where they are better protected from fire and manual damage than those on stumps. Root crowns can ordinarily be killed only through extreme soil disturbance [23]. Root crown buds may not sprout as quickly as stump buds because of the cooler microenvironment at and below the soil surface. Sprouting response is generally slower the deeper a bud is buried [349].

Layering: Baldwin et al. (2012) indicate that salmonberry does not root at the tips [16]. However, Barber (1976) observed several plants on sites near Sedro Woolley, Washington, that had downward-arching stems with tips that were buried by litter. These had rooted and produced new shoots where they came in contact with the ground. Barber also suggests that aerial stems that are damaged mechanically can root and produce new clones when in contact with the ground [18].

Pollination and Breeding System: Salmonberry flowers occur on perennial stems [347]; they are self-incompatible [84] and require cross-pollination. Flowers are pollinated primarily by insects and also hummingbirds [18,189,238,250]. They are also suited to unspecialized pollinators such as beetles [18,238]. Barber (1976) identifies several pollinators of salmonberry flowers [18].

Seed Production: Fruit production depends on environmental conditions, stand characteristics, and salmonberry plant development [238], although large numbers of seeds are typically produced every year [18,238,345,347]. Lepofsky et al. (1985) estimated that salmonberry produced about 30 fruits on a clone covering a 32-foot2 (3-m2) area [188]. The number of seeds per fruit depends on microclimate, pollination, and genetics [347]. Salmonberry averaged 62 seeds/fruit in western Oregon (ranging from 28-128 seeds/fruit) and 40 seeds/fruit in southeastern Alaska (ranging from 17-65 seeds/fruit) [347]. Both red and yellow-orange fruits produce viable seed [238], and samples from Oregon averaged 45 seeds/red fruit and 35 to 42 seeds/orange fruit [319].

Despite abundant seed production, salmonberry seed may not be detected in the seed rain on some sites where it occurs, probably because most fruits are transported offsite by animals (see Seed Dispersal, below). Salmonberry was a common component of the existing vegetation in old-growth western redcedar–western hemlock–Douglas-fir forests (15.9% frequency) and adjacent clearcuts (67.2% frequency) at the University of British Columbia Research Forest near Haney, British Columbia. However, salmonberry seed rain was detected only on clearcuts, and only during 1 of 3 collection years (4 seeds/m2, 1.6% frequency) [159]. Salmonberry seed was not detected in seed rain samples from any site in riparian communities along third- and fifth-order streams on the western slope of the Oregon Cascade Range, although it was present to abundant on most sites [113].

Seed Dispersal and Predation: Salmonberry seed is dispersed mostly by birds and mammals that eat the fruit and deposit the seed in their feces. Passage through a consumer's gut usually enhances germination [318], although digestion by some consumers may kill some seeds (e.g., [90,134]). Salmonberry seed is also dispersed by gravity, when ripe fruits drop from the plant [18,238,345,347]. Seed may be further dispersed throughout the soil by burrowing animals [238], or lost through predation—mostly by rodents [20]. A relatively long period of fruiting in salmonberry increases the probability of seed dispersal by animals [18]. See Importance to Wildlife for more information on animals that consume salmonberry fruit and seeds.

Because dispersal away from the parent plant depends on animals, the number of seeds removed and subsequent distribution of those seeds through space and time depends not only on the size of the fruit, but also on the size, eating habits, and movement habits of the animals. For example, large animals such as grizzly and American black bears are important consumers of salmonberry fruits and are effective seed dispersal agents [318]. Grizzly bears may deposit 50,000 to 100,000 salmonberry seeds in a single pile of feces. Seeds may then be predated or dispersed from feces by small rodents and birds [20,347]. In southeastern Alaska, however, seed predation was less likely for seeds within grizzly bear feces than for clean seeds [20].

Although digestion of fruits typically enhances germination by scarifying seeds, digestion by some animals may reduce viability of salmonberry seeds. Pacific martens may disperse salmonberry seeds long distances (>1,600 feet (500 m)); however, salmonberry seed may not be viable after ingestion by martens [13,133,134]. Banana slugs have been observed eating salmonberry fruits, and they disperse seeds over short distances [90,318]. However, seeds were less likely to germinate after passage through banana slug guts than undigested seed. The negative effects on germination were greater on seeds from red fruits than those from orange fruits [90].

Color preferences of avian frugivores varied among individuals, but most studies found that birds favored red fruits over orange fruits (e.g., [37,89,318]). This may be because the red fruits are mistaken for thimbleberry fruits, which are generally preferred [37]. However, preferences may vary among species of birds and mammals (review by [347]), and one study found that American robins either had no preference or preferred orange fruits over red fruits [340].

Several studies have examined the possible causes or implications of fruit color polymorphism and its possible effects on selection by frugivores. For example, Gervais et al. (1999) reported that birds removed red salmonberry fruits faster than orange fruits in Oregon and Alaska (P<0.01), despite wide geographic variation in fruit-color frequencies, fruit-crop densities, and numbers and species composition of avian frugivores. Based on these results, the authors suggest that forces other than animal selective pressure affect occurrence of salmonberry fruit color [89]. For more details on the interactions and mutualism between avian dispersers and salmonberry fruit color and phenology, see the following publications: [36,37,38,39].

Animals that potentially prey on salmonberry seeds are mostly rodents (squirrels, mice, and voles) and ground-foraging birds. Although red squirrels are important seed predators, they can also disperse seeds by dropping partially eaten fruits. Seed predators did not discriminate between seeds from different colored fruits [318]. Given the large numbers of seeds produced and longevity of seeds in the soil seed bank (see below), seed predation appears to have minimal impact on salmonberry reproduction [238]. In thinned, unthinned, and clearcut conifer stands on two sites in the Oregon Coast Ranges, there was little evidence of salmonberry seed predation [306].

Seed Banking: Salmonberry seeds are dormant when ripe, and form a persistent soil seed bank [18,238]. They can remain viable in the soil for many years or decades [347]. A review by Oleskevich (1996) suggests they remain viable at least 100 years [238]. High annual seed production can result in large seed banks [238]. Estimates of 2 to 125 salmonberry seeds/m2 have been suggested for soil seed bank density [238].

Early observations of abundant salmonberry seedling establishment after logging and soil scarification suggest that abundant viable salmonberry seed occurred in the top 6 inches (15 cm) of mineral soil in second-growth Sitka spruce–western hemlock–Douglas-fir forests in the Oregon Coast Ranges. The author suggested that burrowing moles facilitated salmonberry seed burial on these sites [267].

Several seed banking studies have noted salmonberry emergence from both soil and litter samples on both disturbed and undisturbed sites (e.g., [101,203]), although seed bank density tended to be greater on undisturbed sites (e.g., [203,238]). On some sites, salmonberry was present in the seed bank, despite being absent or infrequent in the aboveground vegetation (e.g., [101,159,203]). Salmonberry tended to be more frequent in the aboveground vegetation on disturbed sites, but it was more frequent in the seed bank of undisturbed sites in British Columbia (e.g., [159,203]). For example, salmonberry was more frequent in the existing vegetation on clearcuts (67.2% frequency) than in adjacent old-growth western redcedar–western hemlock–Douglas-fir forests (15.9% frequency). However, viable salmonberry seed was not detected in the upper 4 inches (10 cm) of surface litter and soil on clearcuts, while 20.3 seeds/m2 emerged from samples taken from old growth [159]. In riparian communities along third- and fifth-order streams on the western slope of the Oregon Cascade Range, salmonberry occurred on most site types, but it was most abundant and dominated the understory on vegetated gravel bars along third-order streams, where it also dominated the soil seed bank, along with stink currant and several herbaceous species. It had low cover (0.1%-0.4%) in old growth dominated by Douglas-fir and western hemlock along both stream types, but it was present in the soil seed bank under old growth along third-order (but not fifth-order) streams [113].

Burial may increase salmonberry seed longevity. On sites in the Sitka spruce-western hemlock zone on the Olympic Peninsula, salmonberry germinants emerged from both soil and litter samples, although germinants were more frequent in soil than litter [101].

Germination: Salmonberry seeds have a dense, impermeable seed coat that contributes to deep dormancy. Dormancy may be broken by a combination of factors including changes in temperature, passage through the digestive system of animals, and activity of fungi and insects on the seedcoat [18,99,347]. Depth of burial, light, and germination substrate (e.g., soil type, feces composition) may affect germination patterns. It is unclear whether seeds from different colored fruits have different germination requirements. Salmonberry seeds from both orange and red fruit showed similar germination patterns in one study [317], and different patterns in another [318]. See Zasada and Tappeiner (2008) for information on germinating salmonberry and other Rubus species under laboratory or greenhouse conditions [347].

Germination patterns vary with microclimate and seed condition when dispersed [347]. Stratification occurs naturally as seeds mature in summer and remain in the soil throughout the cold winter months [238]. Evidence suggests that the action of avian gizzards or exposure to mammalian digestive acids provides beneficial scarification which enhances germination [18,317,318,347]. However, digested seeds receive varying degrees of scarification and may not always remain viable. For example, salmonberry seeds in bear feces may have the fruit wall completely removed or relatively intact [347]. Traveset and Willson (1997) found that passage through vertebrate frugivores enhanced germination of salmonberry seed, with no differences in germination between bear- and bird-ingested seeds. Their results suggest that seed retention time in the guts (much greater in bears than in birds) does not relevantly affect germination [317]. The effect of seed passage through the digestive tract of frugivores (birds and bears) on germination was similar for seeds from both fruit colors, although there were some differences in germination patterns among seeds passed by different frugivores [318].

Most of a given cohort of seeds germinate over a period of 2 to 3 or more years under field conditions, and some seeds apparently lie dormant for decades [347] and germinate when the soil is disturbed. Soil disturbance may be critical for salmonberry germination from the soil seed bank, because germination decreases with increasing depth of burial. Salmonberry consistently emerged on disturbed mineral soils, but it had low emergence rates on undisturbed forest floor [18,238]. Soil scarification after logging results in an excellent seedbed for salmonberry [267].

Salmonberry does not require light to germinate [18,202,347], and germination can occur at relatively low temperatures [18]; however, increased light and temperature (conditions normally associated with soil disturbance) can stimulate germination [238]. Most germination occurs during the first growing season after disturbance. Ruth (1970) observed that germination during the second growing season after overstory thinning was 6% of that which occurred during the first growing season [267].

The type of soil on which seeds are deposited can influence their germination. For example, salmonberry had low germination on alluvial sand and did not germinate at all on reservoir sediments along the Elwha River, Washington [215]. Seeds from one fruit color may germinate better than those from other fruit colors in some soil types. Such differences may contribute to the regional differences in frequencies of the two fruit colors [318].

Seedling Establishment and Plant Growth: Salmonberry seedling establishment is best on bare mineral soil; litter and duff can inhibit establishment [18]. Therefore, seedling establishment is best on disturbed soils [18,268] and is likely to be high after fires that consume surface organic layers (litter and duff). However, seedling establishment is also inhibited by dry conditions, and bare mineral soil can quickly dry, especially in full sun, resulting in high mortality of salmonberry seedlings [238,267]. For example, Ruth (1970) observed up to 500,000 seedlings per acre (1,250,000 seedlings/ha) on scarified soils after overstory thinning but found decreasing numbers of first-year seedlings (although greater overwinter survival) with increasing insolation [267]. Mean salmonberry seedling survival rate was 32% over the first 3 years at four sites in the Oregon Coast Ranges [197].

Because salmonberry seed is commonly dispersed in animal feces, the composition of the feces can affect seedling establishment. Often, feces have a fertilizing effect. However, bear feces usually contain tens of thousands of salmonberry seeds such that emerging seedlings are subject to severe competition and subsequent mortality [318].

Tappeiner and Zasada (1993) studied emergence, survival, and growth of salmonberry on disturbed and undisturbed soil in thinned, unthinned, and clearcut conifer stands on two sites in the Oregon Coast Ranges. Emergence of salmonberry was greater on mineral soil than on soil in which the organic layers were intact. Seedling emergence and survival were greater in thinned stands than in clearcuts or unthinned stands. After 4 years, height of salmonberry was greatest in the clearcuts, where it averaged 9 inches (23 cm) [306].

After establishing from seed in a new site, salmonberry stem number increases rapidly for 1 to 2 years, after which vegetative growth and reproduction result in increased stand density and extensive clonal colonies. Within 2 to 3 years after initial establishment, rhizomes may spread up to ~540 feet2 (50 m2) from the parent plant, and canopy closure may be complete [197,309]. Sprouts arising from buds on aerial stems and rhizomes show greater initial growth rates and greater mean survival rates than seedlings (100% and 70%, respectively); thus, populations are dominated by ramets [238]. Under favorable conditions, height growth can be as much as 5 to 6.5 feet (1.5-2 m) in a single growing season [18,99].

Caplan and Yeakley (2013) examine and compare the relationship between morphological characteristics and growth rates of salmonberry and other Rubus species [43], and Maxwell et al. (1993) provide a model of salmonberry population establishment and growth based on field data from the Oregon Coast Ranges [198].

SUCCESSIONAL STATUS:
Shade tolerance: Salmonberry has relatively high shade tolerance compared to other Rubus species [238], but it does not grow well in deep shade. Seedlings may establish best under moderate shade. Unless soils are moist, seedlings may desiccate and die in areas of high insolation. Established salmonberry plants grow best in full light as long as their roots are in perennially moist soil—the densest salmonberry thickets are in clearings and forest openings rather than under dense overstories [267].

On sites in the central Oregon coast, salmonberry seedlings established better under moderate shade than full sun, and seedlings even tolerated deep shade. Seedling establishment and survival was better under a thinned forest canopy than in a clearcut, and number of seedlings decreased with increasing radiation, indicating that some shade was important for salmonberry seedling survival on these sites. Seedlings that established on open plots did not survive more than 1 year, and no salmonberry seedlings occurred on open plots in the second year. However, little of the variation (6%-16%) was explained by radiation alone. The author suggests that receding soil moisture limited seedling establishment in the open plots more than too much light [267]. Studies in southeastern Alaska compared the response of aboveground growth rates of salmonberry seedlings to simulated variation in light, soil environment [106], and simulated herbivory [105]. Salmonberry growth responded to interactions of canopy and soil (red alder canopy and “mixed” soil producing greatest growth) and to the effects of soil (mixed > mineral) and canopy (red alder canopy > conifer canopy) separately (P < 0.05) [106]. Salmonberry growth and survival were reduced under low light levels and by clipping (P < 0.10) and by low light and clipping combined (P = 0.027), emphasizing the importance of light in determining salmonberry response to herbivory [105].

Salmonberry grows and spreads rapidly after canopy removal and is most abundant on forest sites in early secondary succession, especially on moist soils. In a second-growth Sitka spruce, western hemlock, and Douglas-fir forest on the central Oregon coast, salmonberry cover decreased from 3.1% before overstory thinning, to 1.1% immediately afterward (largely due to mechanical damage), and then consistently increased, reaching 14.6% cover 7 years later [267]. In southwestern British Columbia, increased light availability from partial overstory harvest stimulated salmonberry growth, especially on moist sites [217].

Greater salmonberry biomass in the understory of red alder stands compared with that in conifer stands is likely due to the more open canopy under red alder that allows greater light transmission to the understory [309]. Salmonberry produces leaves earlier in spring than red alder, possibly allowing it to take advantage of the higher light availability before red alder leaves have expanded [18,309].

While it persists in late-successional and old-growth forests in some locations, salmonberry typically declines in abundance as the canopy closes, and it may not persist under a closed canopy. In old-growth forests salmonberry occurs primarily in openings and along edges. In a chronosequence study in southeastern Alaska, for example, salmonberry was most abundant in early succession, up to about 25 to 35 years after removal of the Sitka spruce-western hemlock canopy, declined as tree canopies closed, and increased later in succession (around 140-160 years) as gaps formed in old-growth forests [5].

A comparison of photosynthetic capacity of four endemic plant species in the coastal temperate rainforests of Prince William Sound, Alaska (salmonberry, Sitka alder, Alaska blueberry, oval-leaf huckleberry), showed that differences among species were correlated to their position along the natural light gradient created by the forest canopy [252].

Succession: Salmonberry is an early-seral dominant on moist sites after overstory removal and persists into late forest succession on some sites—mostly in openings and edges. It generally increases in cover after overstory removal, mostly by sprouting from rhizomes, root crowns, and stumps, depending on severity of understory and soil disturbance. Seedling establishment can be very high when disturbance exposes mineral soil (e.g., [5,8,18,268]). See Regeneration Processes for details. Salmonberry is tolerant of frequent flooding that occurs in some riparian communities, and it may form persistent shrub communities in these areas. It may occur in primary succession on some sites.

Salmonberry establishes and spreads in early succession in Sitka spruce-western hemlock forests and in many Douglas-fir forests of the Pacific Northwest [76]. The large number of publications describing its abundance after logging and site preparation provide abundant evidence of its role in early secondary succession (table 5). Salmonberry may establish soon after disturbance by vegetative regeneration or seedling establishment from the soil seed bank (e.g., [8]).

Although most common in early seral stages, salmonberry has also been reported in early immature, second-growth, mature, and old-growth forests in Alaska [5] and British Columbia [170,188]. Percent cover of salmonberry was similar among forest age classes in Douglas-fir stands in western Oregon and Washington (P < 0.05) [287]. In northern California, salmonberry was a common component of the plant community 5 to 10 years after logging in grand fir–Douglas-fir–Sitka spruce and redwood–grand fir communities. Salmonberry persisted in redwood communities, although with reduced abundance, for 30 years or more [351]. In many coniferous stands, parts of salmonberry clones senesce, die, and decay as the overstory canopy closes, but then the clone slowly expands as self-thinning of the conifers occurs [5]. After logging or fire in the coastal Sitka spruce–western hemlock forests of southeastern Alaska, areas with the greatest degree of topsoil disturbance tend to favor salmonberry establishment. It frequently forms a dense canopy with only scattered feather mosses and leaf litter beneath. In areas with less soil disturbance, growth of residual shrubs such as salmonberry increases within 5 years of overstory removal. Understory biomass peaks around 15 to 25 years after logging, with salmonberry, blueberry, and trailing black currant making up >90% of understory production in early succession. As the canopy closes at stand ages of 25 to 35 years, shrubs, graminoids, and forbs are virtually eliminated from the understory. Declines in both standing crop biomass and annual production in the understory are related primarily to the elimination of salmonberry and Alaska blueberry. Bryophytes and ferns dominate understory biomass during the following century. An understory of deciduous shrubs, graminoids, and forbs reestablishes after 140 to 160 years and continues to increase, while bryophyte biomass and tree productivity decline [5].

Salmonberry is a common understory species in old-growth, upland forests in the coastal part of its range, although it may persist in late succession only on moist sites. It occurs in old-growth western hemlock–Sitka spruce forests in southeastern Alaska (e.g., [108,241]), typically in openings, on open streambanks, and in meadows (e.g., [310]). In old-growth Sitka spruce-western hemlock forests on the South Fork Hoh River in Olympic National Park, ungulate herbivory—particularly that of Roosevelt elk—largely determines the distribution and abundance of salmonberry. Salmonberry occurred in "refugia" sites, where it was protected from ungulate herbivory. Refugia were observed most frequently on root mats of fallen trees or behind barriers created by a matrix of fallen trees [272]. Salmonberry is a common component of old-growth western hemlock forests in the northern Oregon Coast Ranges [137]. It averaged 0.1% to 7.7% cover in most old-growth Sitka spruce and western hemlock forest plots on a site in the northern Oregon Coast Ranges [96]. Salmonberry is not consistently present in old-growth redwood forests (e.g., [313]), but it can be a common component or indicator species on relatively wet and riparian sites (e.g., [186,193]). It averaged >20% cover in old-growth redwood–western hemlock and old-growth redwood–red alder forests in southwestern Oregon and northwestern California [193].

Salmonberry is common in red alder communities on both upland and riparian sites and can reportedly persist almost indefinitely in the understory of red alder or mixed hardwood-conifer stands [18]. Upland red alder sites may be early-seral stages of western hemlock forests [131]. Salmonberry was common under 40-year-old red alders in southeastern Alaska [108]. Its aboveground biomass increased with increasing percentage of red alder basal area in second-growth Sitka spruce–western hemlock–western redcedar in southeastern Alaska [107]. Cover and frequency of salmonberry increased with stand age of red alder in a chronosequence of 14-, 24-, and 64-year-old stands along the Hoh River, Olympic National Park, Washington [191]. In the central Oregon Coast Ranges, salmonberry became increasingly more abundant with increasing age of red alder stands, and stands approaching senescence had dense growth of salmonberry, especially those at low elevations [44].

Because few plants can establish and persist under dense salmonberry cover, salmonberry stands may be quite stable, both in the understory of conifer and hardwood stands and in the open (e.g., [217,307,331]). In areas where salmonberry is dominant, elevated microsites—such as old-growth stumps and other coarse woody debris—may provide refugia for species unable to establish and persist under the salmonberry canopy (e.g., salal and red huckleberry) [162]. As red alder becomes senescent, salmonberry and other brushy vegetation may dominate the riparian area [233,354]. In riparian sites periodic flooding can maintain species such as salmonberry, red alder, and stink currant in long-lived disclimax situations [42,131]. Once salmonberry had established after thinning in a second-growth western hemlock–Douglas-fir forest in British Columbia, continued stem recruitment maintained a dense, stable cover in a riparian area and precluded establishment of conifer seedlings. Even intense disturbance (flooding) did not affect the stability of this salmonberry population [217]. See table 4 for additional studies on succession in riparian communities.

Table 4—Studies of secondary succession in riparian forests where salmonberry occurs.
Location Title Source
British Columbia
Vancouver Island Floodplain succession on the west coast of Vancouver Island Clement (1985) [48]
Washington
Olympic National Park Forest succession in relation to river terrace development in Olympic National Park, Washington Fonda (1974) [74]
Oregon
Willamette River Riverscape-level patterns of riparian plant diversity along a successional gradient, Willamette River, Oregon Fierke and Kauffman (2006) [72]
Willamette River Invasive species influence riparian plant diversity along a successional gradient, Willamette River, Oregon Fierke and Kauffman (2006) [71]
Coast Ranges Debris flow through different forest age classes in the central Oregon Coast Range May (2002) [199]
Coast Ranges Ten years of vegetation succession on a debris-flow deposit in Oregon Pabst and Spies (2001) [244]
Coast Ranges Riparian trees, shrubs, and forest regeneration in the coastal mountains of Oregon Minore and Weatherly (1994) [220]

Much information on the role of salmonberry in secondary succession comes from observations following logging. Salmonberry spreads rapidly—from sprouting rhizomes and/or seedling establishment from the soil seed bank—when the overstory is removed and soils are disturbed. Salmonberry dominance in early-seral communities can hinder regeneration and growth of shade-intolerant conifers. In many areas dense stands may form within 2 to 3 years after disturbance. For example, Ruth (1970) observed "almost one-half million seedlings per acre" in the first year after thinning and scarification that removed all ground vegetation but left "as much topsoil as possible" in a 120-year-old Sitka spruce–western hemlock–Douglas-fir stand. Depth of scarification ranged from about 2 to 6 inches (5-15 cm) into the mineral soil [267]. Tappeiner et al. (1991) found that in coastal Oregon forests, the abundance of salmonberry was nearly 300% greater in logged than unlogged stands, and annual rhizome extension averaged 27% in 2- to 3-year-old clearcuts, compared to 2% to 6% in undisturbed stands [264,309]. See table 5 for information on publications that describe succession after logging and site preparation (e.g., scarification, slash burning) and other human-caused disturbances.

Table 5—Publications that provide information on the successional status of salmonberry after human-caused disturbance in different locations throughout its range.
Location Plant Community Disturbance type Title Source
Pacific Northwest
Douglas-fir region Douglas-fir logging/slash burning Vegetative succession following logging in the Douglas-fir region with special reference to fire Isaac (1940) [150]
Alaska
southeastern Alaska Sitka spruce-western hemlock logging Plant succession following logging in the Sitka spruce-western hemlock forests of southeast Alaska Alaback (1984) [6]
British Columbia
coastal British Columbia western hemlock-Sitka spruce logging/slash burning Above- and below-ground vegetation recovery in recently clearcut and burned sites dominated by Gaultheria shallon in coastal British Columbia Messier (1991) [214]
coastal British Columbia roadsides mechanical control Vegetation response to right-of-way clearing procedures in coastal British Columbia McGee (1988) [202]
Washington
western Washington Douglas-fir, western hemlock-Sitka spruce clearcut Six years of plant community development after clearcut harvesting in western Washington Peter (2009) [247]
Oregon
Coast Ranges Douglas-fir thinning Overstory and understory development in thinned and underplanted Oregon Coast Range Douglas-fir stands Chan et al. (2006) [45]
Siuslaw National Forest Douglas-fir clearcut/site preparation Predicting growth response of shrubs to clear-cutting and site preparation in coastal Oregon forests Knowe (1997) [174]
Coast Ranges red alder logging Vegetation characteristics of alder-dominated riparian buffer strips in the Oregon Coast Range Hibbs and Giordano (1996) [132]
Cascade Head Experimental Forest Sitka spruce-western hemlock variable thinning Long-term response of understory vegetation to stand density in Picea-Tsuga forests Alaback (1988) [7]
Coast Ranges red alder chemical control/burning Initial plant succession after brown and burn site preparation on an alder-dominated brushfield in the Oregon Coast Range Roberts (1975) [260]
Siuslaw National Forest Douglas-fir logging The inter-relationship of salmonberry and Douglas-fir in cutover areas Allen (1969) [9]
H. J. Andrews Experimental Forest Douglas-fir clearcut Occurrence of shrubs and herbaceous vegetation after clear cutting old-growth Douglas-fir Yerkes (1960) [344]
California
coastal redwood region redwood-riparian logging The influence of timber harvest on the structure and composition of riparian forests in the coastal redwood region Russell (2009) [265]

Salmonberry may be a component of plant communities in primary succession following deglaciation and volcanic eruption, and on islands (table 6). For example, plant communities on the small islands in Barkley Sound, off the west coast of Vancouver Island, British Columbia, were assembled from seeds dispersed by fruit-eating birds [40].

Table 6—Studies of primary succession in areas where salmonberry occurs.
Location Disturbance type Title Source
Alaska
Glacier Bay Glacial retreat A foundation of ecology rediscovered: 100 years of succession on the William S. Cooper plots in Glacier Bay, Alaska Buma et al. (2017) [33]
British Columbia
Barkley Sound Island assembly Patterns in the assembly of an island plant community Burns (2007) [40]
Washington
Mount St. Helens Volcanic eruption Primary succession trajectories on pumice at Mount St. Helens, Washington del Moral et al. (2012) [62]
Coleman Glacier Glacial retreat Dispersal and establishment both limit colonization during primary succession on a glacier foreland Jones and del Moral (2009) [154]
Coleman Glacier Glacial retreat Patterns of primary succession on the foreland of Coleman Glacier, Washington, USA Jones and del Moral (2005) [153]
Mount St. Helens Volcanic eruption The role of refugia and dispersal in primary succession on Mount St. Helens, Washington Fuller, R. N.; del Moral (2003) [85]
Mount St. Helens Volcanic eruption Ecosystem recovery following a catastrophic disturbance: lessons learned from Mount St. Helens Crisafulli et al. (1998) [52]
Mount St. Helens Volcanic eruption Early primary succession on Mount St. Helens, Washington, USA del Moral et al. (1995) [61]
Mount St. Helens Volcanic eruption Plant form, developmental plasticity and survival following burial by volcanic tephra Antos and Zobel (1985) [11]
Mount St. Helens Volcanic eruption Natural revegetation of the northeastern portion of the devastated area Means et al. (1982) [208]

FIRE EFFECTS AND MANAGEMENT

SPECIES: Rubus spectabilis
FIRE EFFECTS
Immediate Fire Effects
Salmonberry is moderately to highly resistant to fire [18,238,332]. Rhizomes and root crowns generally survive even when all aerial stems are killed. Therefore, actual plant mortality from fire is presumably low. Seed in the soil seed bank is probably unharmed by fire, depending on depth of burial, while that in the litter and duff may be consumed or killed by severe fire that consumes surface organic layers.

Postfire Regeneration Strategy [297]
Tall shrub, adventitious buds and/or a sprouting root crown
Rhizomatous shrub, rhizome in soil
Geophyte, growing points deep in soil
Ground residual colonizer (on site, initial community)
Initial off-site colonizer (off site, initial community)
Secondary colonizer (on- or off-site seed sources)

Fire Adaptations and Plant Response to Fire
Fire Adaptations: Salmonberry reproductive traits (i.e., buried seed, buried rhizomes, and rapid sprouting following top-kill) make it moderately to highly resistant to fire [238]. Stephens et al. (2018) classify salmonberry as a "fire neutral, facultative sprouter", meaning that individual plants are top-killed, but roots and rhizomes generally survive fire. Top-kill from fire stimulates sprouting from buds on rhizomes, root crowns, and stumps; and fire creates a good seedbed for seedling establishment—primarily from the soil seed bank (when present) and secondarily from animal-dispersed seed [292]. High-severity fires (i.e., with high surface fuel consumption and severe soil heating) may reduce salmonberry cover in the short term (~10 years), especially on dry sites, by killing buried seeds and shallow buds on root crowns and rhizomes, reducing postfire seedling establishment and sprouting [99,238].

Plant Response to Fire: Salmonberry is a common component of the postfire environment in coastal forests and shrublands and an early-seral dominant, especially on moist sites [2,25,31,47,181]. It is likely to increase in abundance and productivity (fruiting) after fire [205,238,246]. Severe fire—such as that from slashburning—may limit or slow salmonberry recovery, although the effects of slashburning vary.

Dense stands of salmonberry can develop within 2 to 5 years after fire [127,308], and salmonberry remains abundant in communities on moist sites until the canopy closes [308]. For example, in Engelmann spruce forests of the upper Fraser River valley in central British Columbia, salmonberry occurred in stands 4 to 22 years after fire, but it was not recorded in stands 37 to 75 years after fire [86].

On the Tillamook Burn in northwestern Oregon, postfire plant communities reflected their prefire distribution. Three successive, high-severity (i.e., stand-replacement) fires (1933, 1939, and 1945) in the Tillamook Burn in the northern Oregon Coast Ranges killed the conifers and left vegetation dominated by pioneer species including red alder, vine maple, western brackenfern, salal, salmonberry, and other Rubus spp. over an area of approximately 300,000 acres (~121,000 ha) [135,229]. On burned sites, salmonberry dominated the understory, along with thimbleberry, western swordfern, and Siberian springbeauty, beneath a canopy of red alder and bigleaf maple [14]. One block of unburned forest, approximately 2,500 acres (~1,000 ha) in size, remained in the center of the burn; it was estimated at 300 years old and had no evidence of prior disturbance. Salmonberry was present in the undisturbed forest, but at low frequency (1%); it was more frequent (12%) in the adjacent burned area, which had also been salvage logged [229].

On some upland sites in the Pacific Coast Maritime region, dense shrub communities dominated by salmonberry, salal, red huckleberry, and vine maple develop after stand-replacing fires [2,312]. Postfire shrubfields may persist indefinitely or eventually be replaced by conifers that regenerate beneath the shrub canopy [312]. In northern Humboldt County, California, thickets of salmonberry and thimbleberry established after several prescribed burns; the thickets were sometimes interspersed with thickets of Pacific poison-oak [35]. Near Blaney Lake, British Columbia, in a forest dominated by western redcedar, western hemlock, and Douglas-fir, a severe fire in the mid-1800s was followed by clearcutting and slashburning in the 1920s and another fire in 1931. In depressions where the soil was damp but lacked standing water in summer, shrubby species formed dense thickets that occupied a considerable area of the burn. The most common species was salmonberry, with thimbleberry and rose spirea less common. Salmonberry was 2 to 6 feet (0.6-1.8 m) tall, and its height and abundance varied with soil moisture. Salmonberry also grew under tall western brackenfern in areas where western brackenfern was dominant. Salmonberry was a minor component in the predisturbance forest, occurring at low densities in damp depressions, edges, and openings [205].

Vegetative response: Salmonberry sprouts following top-kill by fire [238]. Overstory mortality, fire intensity, and soil burn severity influence the rate of salmonberry postfire recovery [192]. When stumps survive, stump-sprouting is the predominant mode of postfire regeneration. Apical dominance suppresses sprouting from root crowns and rhizomes [349]. The root crown has some protection from the direct effects of fire because it is located at or below the soil surface, and buds on the root crown may sprout after top-kill if buds on the stump are killed. Root crown buds are generally killed only by extreme soil disturbance. If the root crown is killed, underground rhizomes typically sprout prolifically. Rhizomes are protected from fire by overlying soil, and their extensiveness insures that some are likely to survive. Even an extremely severe fire is unlikely to kill the entire network of well-protected rhizomes. Aerial stem growth from buds on root crowns and rhizomes tends to be somewhat slower than from stumps because of the cool underground environment in which these sprouts develop [349].

Salmonberry plant age and condition, which is related to site characteristics, can affect postfire response. Younger rhizomes tend to grow more actively and sprout more vigorously than larger, older rhizomes, which typically have lower bud densities [349]. Rhizomes tend to be most developed on relatively mesic sites with deep soils, and they may be poorly developed or even lacking on dry, rocky sites [308]. Postfire sprouting could presumably be reduced on sites with shallow or rocky soils, or where the prefire stand was primarily made up of older plants lacking vigor.

Fire timing may affect postfire response of salmonberry. Salmonberry seems to have sufficient carbohydrate stores to sprout after a single fire regardless of timing. However, seasonal changes in TNC content in rhizomes may affect the number of shoots produced after top-kill from fire. Low shoot production coincides with low levels of TNC, which occur during spring shoot production and early summer growth. Therefore, salmonberry regrowth may be slower after fires during the growing season (May through July) [291,348].

Seedling establishment: Salmonberry seeds may remain viable for years when buried in the soil or duff [18], and this soil seed bank is an important mode of postfire establishment. Seedlings require mineral soil for best establishment and growth, so when fire consumes litter and duff and exposes mineral soil, a good seedbed is present for salmonberry seedling establishment [238]. Some seedling establishment can also occur through seed transported from off site (see Seed Dispersal).

Response to Slashburning: Salmonberry response to slashburning depends on its physiological conditions (age and stem density), site characteristics, and overall fire severity (i.e., depth and duration of soil heating). Low- to moderate-severity slash fires generally increase salmonberry cover compared to unburned controls [99]. High-severity slash fires, particularly on dry sites, may reduce salmonberry cover in the short term (~10 years) but can also cause site degradation from severe soil heating [99,238]. In some instances, particularly where salmonberry was abundant before logging, salmonberry grows rapidly despite severe slash fires because only a small portion of the extensive rhizome network is damaged (e.g., [288,349]). Zasada et al. (1989) cite an example where stem growth of salmonberry was rapid, and pretreatment height was reached by mid- to late summer on sites logged and burned in February, March, and April; whereas pretreatment height was not reached until the end of the growing season on sites that were logged and not burned [349]. For additional studies that examine plant community response to slash burning in areas where salmonberry occurs, see table 7.

Table 7—Publications with information on salmonberry response to logging and burning.
Location Plant Community Disturbance type Title Source
Pacific Northwest
Washington, Oregon Douglas-fir slash burning Effects of slash burning in overmature stands of the Douglas-fir region Morris (1970) [223]
Washington, Oregon Douglas-fir slash burning Influence of slash burning on regeneration, other plant cover, and fire hazard in the Douglas-fir region: a progress report Morris (1958) [222]
Washington, Oregon Douglas-fir logging and burning Vegetative succession following logging in the Douglas-fir region with special reference to fire Isaac (1940) [150]
Alaska
southeastern Alaska Sitka spruce–western hemlock slash burning Effects of slash burning on conifer regeneration in southeast Alaska Harris (1966) [119]
British Columbia
northern Vancouver Island Douglas-fir–western hemlock–western redcedar clearcut, slash burn Above- and below-ground vegetation recovery in recently clearcut and burned sites dominated by Gaultheria shallon in coastal British Columbia Messier and Kimmins (1991) [214]
British Columbia forests slash burning Site preparation: fire Hawkes et al. (1990) [127]
western Vancouver Island western redcedar–western hemlock clearcut, slash burn Effects of prescribed burning on vegetation and natural tree regeneration in mature cedar-hemlock forests in the Pacific Rim National Park McGeough (1985) [204]
Vancouver Forest Region forests slash burning Site diagnosis, tree species selection, and slashburning guidelines for the Vancouver Forest Region [97,168]
Vancouver Island Douglas-fir slash burning Some effects of slash burning on regeneration and growth of Douglas-fir (Pseudotsuga menziesii) on Vancouver Island, British Columbia Knight (1961) [173]
Oregon
Coast Ranges Douglas-fir–western hemlock–western redcedar logging and burning Forest associations and secondary succession in the southern Oregon Coast Range Bailey (1966) [15]
Coast Ranges Douglas-fir clearcut, spraying, and burning Origin and development of vegetation after spraying and burning in a coastal Oregon clearcut Stewart (1978) [296]

FUELS AND FIRE REGIMES
Fuels: No information specifically describing salmonberry fuel characteristics was found in the available literature as of 2019. Some studies are available that provide information on salmonberry biomass and stand structure in particular site types. For example, Hanley et al. (2006) provide data on salmonberry leaf, twig, and stem biomass under red alder stands with varying basal area in southeastern Alaska [107], Tappeiner et al. (1991) provide data on salmonberry stand structure and biomass in different plant communities in western Oregon [309], and Van Pelt et al. (2016) give data regarding leaf area and biomass of salmonberry in California redwood forests [328].

Fuel guides that include vegetation associations where salmonberry is an understory dominant are available (e.g., [221,249]). Also see the Natural Fuels Photo Series and Digital Photo Series websites.

Fire Regimes: Fires were historically infrequent in many of the coastal forests where salmonberry occurs (e.g., [181]). For example, Sitka spruce–western hemlock forests were thought to have long fire-free intervals (~300-1,000 years), and fires were thought to be stand replacing when they did occur. Windthrow is a more common disturbance in these forests [181]. Salmonberry occurs in forests that are transitional between coastal Sitka spruce–western hemlock forests and white spruce–paper birch boreal forests of south-central Alaska [226]. Coastal forests in Alaska rarely, if ever, burned historically [355], while white spruce-dominated boreal forests had long intervals (~100-800 years) between high- and mixed-severity fires [1]. Salmonberry also occurs in plant communities with short to moderate historical fire intervals, such as those dominated by redwood, Douglas-fir, and western larch. Stephens et al. (2018) provide a summary of historical fire regimes in redwood forests [292].

Salmonberry occurs in plant communities where lightning fires were historically rare or infrequent, largely due to the rarity of lightning, but also due to the wet climate. For example, lightning is rare in the coastal mountain ranges where salmonberry thrives. However, in those areas fires were regularly ignited by American Indians for a variety of purposes, including but not limited to creating and maintaining patches of fruit-bearing plants, such as salmonberry, and improving their yield (e.g., [24,26,65,180,292]). In addition, the berry patches created by regular burning were also excellent habitat for game animals. For example, elk and deer prefer to browse on young plants growing in burned areas, including the leaves and stems of salmonberry [24]; see Importance to Wildlife for additional information. A burning schedule with 3-year intervals was said to be best for berry production in general. Burning may have been conducted in either spring or fall [187], but often occurred after the berry harvest (e.g., [24,26]). No berries would occur in the burned patch the year after burning, but in 2 years, the berry crop would be larger than before [24].

Because American Indians actively managed salmonberry with fire, areas where salmonberry patches are widespread may have a history of regular fire. Sometimes fires escaped, and berry patches became much larger than intended [24]. Landscape vegetation patterns seem to support this. Salmonberry patches along small rivers and streams in the Oregon Coast Ranges are an example of local-scale patterns of managed vegetation that may have persisted from presettlement times. While shrubs (such as salmonberry and huckleberry), forbs, and perennial grasses are not as reliable for interpreting fire pattern as are trees, they do form identifiable landscape patterns that can persist decades or centuries. Many areas where conifer establishment was prevented by regular burning before Euro-American settlement remained without conifer forests at the turn of the 21st century [356]. This is not the case in all areas. For example, in the absence of fire, areas of former coastal prairie that were once maintained by regular, anthropogenic burning are now dominated by forest species including Sitka spruce, redwood, western hemlock, red alder, Cascara buckthorn, salmonberry, and salal [180].

When fires did occur in coastal forests, they likely covered large areas with high-severity effects (i.e., stand-replacement). Large and high-severity fires were more likely to occur in the wetter landforms, vegetation types, and geographic positions (i.e., the western and central part) of the central Oregon Coast Ranges than in drier sites because fires were less frequent, allowing greater fuel accumulation. The higher fuel loads would then facilitate fire spread across the landscape and into the canopy when weather conditions were suitable for drying fuels and spreading flames [149].

Little is known about presettlement fire regimes in plant communities where salmonberry occurs in northern coastal scrub in the north coast bioregion of California, although fire is thought to have been relatively uncommon. Northern coastal scrub is self-perpetuating with or without fire. Fires that did occur historically were most likely ignited by American Indians, and fire intervals may have ranged anywhere from 1 to 100 years, depending on vegetation composition and proximity to human settlements [292].

Because many of the plant communities in which salmonberry occurs are characterized by infrequent fires, a longer term perspective on fire regime variability may be desirable and informative. Fire history studies that employ methods to analyze fire frequency over millennia (e.g., dating charcoal deposits in lake sediments) include one from the Oregon Coast Ranges [190] and two from Vancouver Island [32,88].

More information on fire regimes in plant communities where salmonberry occurs is available in these literature reviews [2,292] and in the FEIS Fire Regime publications shown in table 8.

Table 8—FEIS publications with information on fire regimes in ecosystems where salmonberry is a common to dominant species in one or more Biophysical Settings.
Region Title
Alaska Fire regimes of Alaskan Pacific maritime ecosystems
Alaska Fire regimes of Alaskan mountain hemlock ecosystems
Alaska Fire regimes of Alaskan alder and willow shrublands
Alaska Fire regimes of Alaskan black cottonwood communities
Pacific Northwest Fire regimes of Pacific Northwest riparian communities
Pacific Northwest Fire regimes of North Pacific montane shrubland communities
Pacific Northwest Fire regimes of Pacific Northwest coastal forests
Pacific Northwest Fire regimes of red alder landslide communities
Pacific Northwest Fire regimes of wet-mesic western hemlock communities
Pacific Northwest Fire regimes of Pacific Northwest wooded volcanic flowage communities
California, Pacific Northwest Fire regimes of montane riparian communities in California and southwestern Oregon
California, Pacific Northwest Fire regimes of coastal scrub communities
California, Pacific Northwest Fire regimes of redwood communities
California, Pacific Northwest Fire regimes of mediterranean mixed-evergreen communities

FIRE MANAGEMENT CONSIDERATIONS

Salmonberry provides food and cover for many wildlife species, but it may also have negative impacts on wildlife habitat when it persists and spreads. Therefore, the impacts of both fire and fire exclusion are important considerations for wildlife managers of plant communities where it occurs. Because salmonberry can interfere with conifer establishment—especially after logging—the effects of slash burning on its regeneration and abundance have been the subject of much research. See table 7 for references on salmonberry response to slash burning, and see Other Management Considerations for information and references on salmonberry response to logging and other site preparations.

Fire generally benefits animals that consume the fruits of Rubus spp. [177], because fire increases fruit production (e.g., [205,238,246]). Fire may also benefit ungulates that browse on salmonberry by increasing the abundance of new shoots, which are preferred (see Importance to Wildlife).

Fire exclusion may increase salmonberry cover in some plant communities and may have negative impacts on wildlife that use those communities. For example, habitat of the Oregon silverspot butterfly has been reduced in some areas by the spread of salmonberry into early successional grasslands along the Pacific Coast from southern Washington to northern California. Preventing or removing this sort of encroachment by shrubs to maintain or improve grasslands as suitable habitat can be achieved using prescribed fire. Season of burning is important because burning in early fall can kill eggs and larvae of the Oregon silverspot butterfly. In addition, undesirable plants such as tansy ragwort and western brackenfern may increase after fire [248]. Under some conditions, salmonberry cover may also increase after fire (see Vegetative response), so caution is warranted.

MANAGEMENT CONSIDERATIONS

SPECIES: Rubus spectabilis
FEDERAL LEGAL STATUS
None

OTHER STATUS
Information on state- and province-level protection status of plants in the United States and Canada is available at NatureServe. Salmonberry is ranked as "imperiled" (S2) in Idaho [227].

IMPORTANCE TO WILDLIFE AND LIVESTOCK
Numerous wildlife species use salmonberry. A variety of animals eat its fruits, seeds, stems, and leaves, and many birds and small mammals use it for cover [18,102,189], especially in riparian areas (e.g., [78]). For example, songbirds, gallinaceous birds, bears, and coyote eat salmonberry fruit [192]; mice and other small rodents consume salmonberry seeds [18]; and deer, elk, mountain goats, and moose browse on its buds and twigs [192]. Traveset and Willson (1998) provide a list of animals that eat salmonberry fruits in southeast Alaska [318]. Nectar from salmonberry flowers provides food for bees and other insects, as well as for rufous hummingbirds [18,250,298].

Cattle seldom eat salmonberry, but it is considered fair browse for domestic sheep in parts of west-central Washington [58]. In some areas (e.g., Douglas-fir plantations), it is a preferred summer browse for domestic sheep [184], which have been used to control unwanted woody vegetation including salmonberry in regenerating conifer plantations [185,278]. Aerial parts of salmonberry have been used as food for livestock in British Columbia [182].

Fruits and seeds: Several birds eat salmonberry fruits, including American robin, gray catbird [18], varied thrush [19], and band-tailed pigeon [91,144,228]. In addition, salmonberry fruits are important forage for coyote, American black bear, and grizzly bear [18,103,141,147,330]. Banana slugs have also been observed eating salmonberry fruits [90,318]. Small rodents eat salmonberry seeds [18].

Stems and leaves: The foliage, stems, cambium, and bark of species within the Rubus genus provide food for small mammals such as rabbits, North American porcupine, and American beaver [50,327]. Mountain beaver commonly browse salmonberry during the summer [70,102]. It was not an important component in the diet of white-footed voles in Oregon, despite its dominance on study sites [333].

Salmonberry shoots and leaves are important forage for large mammals such as grizzly bears (e.g., [103,147]), American black bears (e.g., [141]), and several ungulates, including deer, elk, moose, and mountain goats. Deer, moose, and mountain goats browse new leaves and twigs in the spring [330]. The tender, leafy new growth is a preferred deer food in many areas [161]. Deer and elk use is often heavy during summer [102]. In Douglas-fir forests in the Oregon Coast Ranges, deer continue to feed on salmonberry leaves until they drop from the plants in autumn [102]. Additional information on use of salmonberry by ungulates is available (table 9).

Salmonberry is an important elk browse in many areas, including the Olympic Peninsula [18,189]. Elk use the leaves and twigs to some extent year-round [273], but use tends to be particularly heavy during the spring and summer [80,102,114,273]. Salmonberry was a common component in elk diets in both old-growth and 1- to 35-year-old second-growth Douglas-fir–western hemlock forests in the Cascade Range, Washington. Greatest use of salmonberry was in early spring and late fall [152]. It was among the shrub and forb species dominating the summer diet of Roosevelt elk in the Mount St. Helens blast zone 5 years after the eruption [211]. Shrub use by Roosevelt elk peaked in September in the blast zone, but comprised >20% of feces sampled during all months (June through November). Willows, red elderberry, blue elderberry, salmonberry, and huckleberry consistently made up most of the shrubs consumed [212].

In Sitka spruce–western hemlock forests of the Pacific Northwest, Roosevelt elk greatly influence the composition and structure of the forest understory by grazing and browsing. Preferred species such as salmonberry are found only in protected microsites [79]. A long-term ungulate exclosure study in Olympic National Park showed greater abundance of salmonberry within exclosures [343].

Mountain goats and moose may browse the young stem tips of salmonberry early in the season [18,192,330]. Only light browsing of salmonberry by mountain goats was observed in subalpine habitats in south-central Alaska in winter [140].

Table 9—Studies on habitat use by ungulates in areas where salmonberry is a common to dominant species.
Location Animal(s) studied Title Source
Pacific Northwest
Alaska to California Roosevelt elk Food habits of Roosevelt elk Jenkins and Starkey (1991) [151]
Alaska
southeastern Alaska Sitka black-tailed deer Relationships between Sitka black-tailed deer and their habitat Hanley (1984) [110]
southeastern Alaska Sitka black-tailed deer, mule deer Response of deer to secondary forest succession in southeast Alaska Wallmo and Schoen (1980) [336]
British Columbia
southwestern British Columbia black-tailed deer Ecology of black-tailed deer Bunnell (1990) [34]
southern Vancouver Island Columbian black-tailed deer 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 Cowan (1945) [51]
Washington
Mount St. Helens elk Population dynamics and habitat ecology of elk in the Mount St. Helens blast zone Merrill et al. (1987) [213]
Columbian White-tailed Deer National Wildlife Refuge Columbian white-tailed deer Habitat use by Columbian white-tailed deer Suring and Vohs (1979) [302]
western Washington black-tailed deer The black-tailed deer of western Washington Brown (1961) [29]
Oregon
Coast Ranges black-tailed deer Black-tailed deer and Douglas-fir regeneration in the Coast Range of Oregon Hines and Land (1974) [136]
Coast Ranges Roosevelt elk Roosevelt elk-forest relationships in the Douglas-fir region of the southern Oregon Coast Range Swanson (1970) [303]
Tillamook Burn, northwestern Oregon Columbian black-tailed deer Forage availability in relation to browsing of Douglas-fir seedlings by black-tailed deer Crouch (1968) [54]
Tillamook Burn, northwestern Oregon Columbian black-tailed deer Food preferences of the Columbian black-tailed deer Odocoileus hemionus columbianus (Richardson) on the Tillamook Burn, Oregon Chatelain (1947) [46]
Tillamook Burn, northwestern Oregon black-tailed deer Management of black-tailed deer Einarsen (1946) [68]
California
Redwoods State Park and surrounding area Roosevelt elk The status and ecology of the Roosevelt elk in California Harper et al. (1967) [115]

Palatability and Nutritional Value: The fruit of salmonberry is highly palatable to many species of birds and mammals, including humans [18]. Salmonberries are described as deliciously flavored, although somewhat variable in taste [58,102,329]. Salmonberry leaves and twigs are reportedly highly palatable to elk from spring through fall [273]. However, the leafy new growth of salmonberry appears to be more palatable to most wild ungulates than the tougher mature foliage [7,58,330]. Dormant twigs are rarely eaten [136] and are presumably somewhat less palatable than those of many shrub associates.

Nutritional value of salmonberry fruits and browse varies both seasonally and annually. For example, crude protein values may be higher in the spring or early summer than in winter [67]. Lipid and sugar content suggest that salmonberry fruits are among those with the highest relative energy reward of fruiting species in southeast Alaska [319]. Table 10 shows additional studies that provide information on salmonberry nutrient composition.

Table 10—Studies providing information on nutritional composition of salmonberry.
Location Information provided Citation(s)
Alaska, Washington, British Columbia Salmonberry nutrient composition Norton (1981) [235], Norton et al. (1984) [234]
Alaska and Washington A breakdown of chemical constituents of salmonberry and other associated forage species used by mule deer and black-tailed deer McArthur et al. (1993) [200]
southeastern Alaska Seasonal changes in chemical composition and nutritive values of salmonberry and other native forages in Sitka spruce–western hemlock forests Hanley and McKendrick (1983) [109]
Olympic Peninsula, Washington Nutritional quality and tannin astringency of salmonberry and other plants browsed by black-tailed deer and Roosevelt elk in clear-cuts and old-growth forests Happe et al. (1990) [112]
northwestern portion of the Mount St Helens blast zone In vitro dry matter digestibility and percent nitrogen content of new twigs and leaves of salmonberry, July through September Merrill et al. (1995) [212]
Oregon Coast Ranges Crude protein values of salmonberry foliage recorded during two seasons Rhodes and Sharrow (1983) [256], Rhodes and Sharrow (1990) [257]

Cover Value: Salmonberry provides cover for birds and small animals, such as mountain beaver [102]. It was commonly dominant in the shrub stratum of ruffed grouse drumming sites in western Washington. Salmonberry has no leaves during most of the drumming season and therefore does not substantially obstruct visibility [270]. It is a common component of streamside vegetation in the Pacific Northwest, where it enhances fish habitat by providing shade and helping prevent or reduce soil erosion [30].

VALUE FOR REHABILITATION OF DISTURBED SITES
Salmonberry is used for rehabilitation of many types of disturbed sites (e.g., [283]). Its deep rhizome and root system can help prevent or reduce soil erosion [18,210].

Salmonberry seedlings and cuttings can be transplanted onto disturbed sites with good results. Salmonberry can be established from root crown or rhizome cuttings under field conditions in coastal Oregon with little post-planting care if planted during the wet season in the winter [197,347]. Cultivars are available [293]. The NRCS's online Plant Guide (2000) has information on establishing salmonberry from cuttings and seeds. See Barber (1976) [18] and Maxwell (1990) [197] for additional information on establishing salmonberry from cuttings, and see Zasada and Tappeiner (2008) for information on collecting, extracting, storing, and germinating salmonberry seeds [347].

OTHER USES
Salmonberry is a traditional food and medicinal plant of American Indians and First Nations peoples along the Pacific Coast (e.g., [18,24,156,188,234,235,236,254,320,322,353]). It is still a very important wild food and medicinal plant in the maritime region of Alaska (e.g., [146,235]), and the fruits may be a potential income source as a nontraditional forest product [321].

People eat young shoots of salmonberry in the spring when little other wild food is available [18,24,102]. The bark and leaves are used to make various medicinal preparations [24,102,182,188,321].

Salmonberry fruits are one of the earliest to ripen (May-June). The ripening is associated with the song of Swainson's thrush, which is called "salmonberry bird" in many languages [251]. Fruits can be eaten fresh or preserved but are rarely dried [58,102,188,320,329].

The plants are ornamental and often cultivated (e.g., [27]) but can become weedy and difficult to eradicate [139].

OTHER MANAGEMENT CONSIDERATIONS
Management of plant communities where salmonberry is a common or dominant species includes considerations for wildlife, nonnative invasive plants, insects and disease, and climate change. Much information is available regarding salmonberry response to logging and site preparation, including means to reduce its cover using a variety of control methods.

Management for Wildlife: Salmonberry provides important habitat for many bird species, therefore disturbances that reduce salmonberry cover may have adverse effects on bird populations. For example, a study conducted in Haida Gwaii suggests that overabundance of Sitka black-tailed deer can decrease songbird habitat quality by decreasing cover of shrubs such as salmonberry, which provide food and nest sites. In areas with severe browsing, the shrub understory—dominated by salal, red huckleberry, and salmonberry—was replaced by an open understory with a ground layer of mosses, reducing both food and cover for birds. Songbird abundance was 55% to 70% lower on islands browsed by deer for more than 50 years than on islands without deer, and alpha diversity was lower on islands browsed by deer. Bird species dependent on understory vegetation were most affected [10]. Pearson and Manuwal (2001) recommend a minimum buffer of 148 feet (45 m) in forested riparian areas along both sides of second- and third-order streams during logging operations to maintain berry-producing shrubs (especially salmonberry) and deciduous trees such as red alder and bigleaf maple, which support breeding bird populations associated with riparian habitats in Douglas-fir forests of the coastal Northwest [245].

Salmonberry can reduce habitat for some birds. For example, Cassin's auklets tend to avoid salmonberry and prefer tufted hairgrass for nesting (e.g., [138,261]). Declines in Cassin's auklet burrow density and population size from 1989 to 2009 on Triangle Island, British Columbia, were associated with increases in salmonberry and decreases in tufted hairgrass cover. Soil disturbance by these burrow-nesting seabirds negatively affected and reduced percent cover of salmonberry, but only when burrow densities were high or increasing. At lower or declining burrow densities, seabird activities were inadequate to halt succession to salmonberry dominance on this nonforested island [261].

Additional studies that examined habitat use by birds in areas where salmonberry is a common component of the vegetation include a comparison of diurnal bird communities among several coniferous and deciduous forest types in Alaska and adjacent Canada [341]; an examination of bird community structure on early-growth clearcuts in western Oregon [224]; and a comparison of bird assemblages in riparian and upland habitats in old-growth coastal western hemlock forest on western Vancouver Island, British Columbia [282].

Large and small mammals that use salmonberry for food and cover may be adversely affected by logging operations. In British Columbia, forest silvicultural practices were altered in coastal riparian areas to provide for adequate fruit production by salmonberry and other species that are important food sources for grizzly bear (review by [347]). Studies that examine the effects of logging and site preparation on small mammals in habitats where salmonberry occurs in Oregon [142,143], Alaska [121], and British Columbia [299] are available.

Site preparations aimed at removing slash and reducing tree seedling interference from shrubs such as salmonberry can have adverse impacts on stream habitat. For example, removing slash and other debris from stream channels after logging by burning or other means can have detrimental effects on fish habitat due to increased water temperatures and soil erosion, and decreased abundance of insects [30].

Nonnative Invasive Plants: Nonnative invasive plants are a common management concern in all wildlands of the Pacific Northwest (e.g., [129]). Nonnative knotweeds are of particular concern in communities where salmonberry occurs. Giant knotweed is invasive in riparian corridors in the Pacific Northwest, where it can reduce richness and abundance of native plants and the quantity and quality of litter inputs [325]. Information on controlling Bohemian knotweed and other invasive plants (Himalayan blackberry, Scotch broom, and reed canarygrass) in western Washington is provided by Danvenport (2006). Part of the control strategy involves planting competitive native species such as salmonberry [56].

Insects and Disease: Ellis et al. (1997) provide an overview of insects and diseases that affect salmonberry and other Rubus spp. [69]. Salmonberry is a host for the pathogen that causes sudden oak death [259].

Management Under a Changing Climate: Ecosystems where salmonberry occurs include those in which the impacts of global climate change have been evident for decades. For example, Alaskan coastal forests have shown reduced frequency of snow avalanches at low and moderate elevations due to more winter precipitation falling as rain and less as snow. One result of this change is that salmonberry is spreading into meadows formerly dominated by mountain heather and sedges [155]. More information on the impacts of climate change in southeast Alaska is available (e.g., [160,355]).

Silvicultural Considerations: Portions of both rhizomes and aboveground stems of salmonberry damaged during silvicultural operations can sprout [18], and salmonberry seedling establishment can be dense in areas with soil disturbance (e.g., [7]). This often results in dense and persistent stands of salmonberry after clearcutting and other types of logging and thinning [7,117,258,264,267,295,315,330]. Brushfields dominated by salmonberry and other shrubs are particularly common at low to middle elevations on moist slopes in the Coast Ranges of Washington and Oregon [93,95,158,179] and on moist valley bottoms in Alaska [330].

Salmonberry brushfields interfere with establishment of conifer and other seedlings [4,9,102,161,184,258,266,268,295,305,323,330]. Dominance of salmonberry in cut-over forestland in the Oregon Coast Ranges likely contributes to a reduction in floral diversity in these ecosystems [207]. Opalach et al. (1990) provide a simulation model of Douglas-fir and salmonberry competition [239,240]. Additional information on salmonberry response to silvicultural operations and its impacts on tree regeneration is available in the publications shown in table 11.

Table 11—Publications with information on salmonberry response to silvicultural operations and/or its impacts on tree regeneration.
Location Plant community Disturbance type Title/topic Source
Pacific Northwest
Washington and Oregon multiple utility rights-of-way Identifying plant communities resistant to conifer establishment along utility rights-of-way in Washington and Oregon, U.S. Shatford et al. (2003) [279]
Douglas-fir region Douglas-fir silvicultural treatments Silviculture for multiple objectives in the Douglas-fir region Curtis et al. (1998) [55]
Coast Ranges Douglas-fir clearcut, site preparation Ten-year development of Douglas-fir and associated vegetation after different site preparation on Coast Range clearcuts Stein (1995) [289]
Pacific Northwest Douglas-fir, western hemlock, Sitka spruce logging Effect of salmonberry on growth of planted conifers Newton and White (1983) [231]
Pacific Northwest forests logging A comparison of harvesting methods and their impact on soils and environment in the Pacific Northwest Cromack et al. (1979) [53]
Pacific Northwest western hemlock logging Harvest cuttings and regeneration in young-growth western hemlock Ruth (1976) [269]
Cascade Range western hemlock–Sitka spruce logging Regeneration and growth of west-side mixed conifers Ruth (1974) [268]
Alaska
southeastern Alaska western hemlock–Sitka spruce logging and site preparation The forest ecosystem of southeast Alaska: Forest ecology and timber management Harris and Farr (1974) [126]
southeastern Alaska western hemlock–Sitka spruce clearcut Natural reforestation on a mile-square clearcut in southeast Alaska Harris (1967) [120]
British Columbia
British Columbia Douglas-fir, western hemlock–Sitka spruce silvicultural treatments A preliminary guide to the response of major species of competing vegetation to silvicultural treatments Coates and Haeussler (1986) [49]
southwestern British Columbia Douglas-fir silvicultural operations Ecology and silviculture of the most productive ecosystems for growth of Douglas-fir in southwestern British Columbia Klinka and Carter (1980) [164]
Washington
Washington Douglas-fir logging, thinning Effects of forest management on understory and overstory vegetation: a retrospective study Thysell and Carey (2000) [314]
Washington western hemlock shelterwood cutting Results of shelterwood cutting in western hemlock Williamson and Ruth (1976) [339]
Oregon and California
Oregon coast Douglas-fir repeated thinning Understory tree development with repeated stand density treatments in coastal Douglas-fir forests of Oregon Shatford et al. (2009) [280]
southwestern Oregon Douglas-fir logging Brush problems in southwestern Oregon Gratkowski (1961) [92]
southwestern Oregon and northwestern California multiple logging, thinning, site preparation Ecology of hardwoods, shrubs, and herbaceous vegetation: effects on conifer regeneration Tappeiner et al. (1992) [304]

Control of Salmonberry: Because it spreads rapidly and forms dense stands after overstory removal, salmonberry is considered a weed in some management contexts (i.e., after logging) [311], and much has been written on methods for controlling it (e.g., [94,230]). Because of its extensive rhizome network, salmonberry is difficult to remove once established. When top-killed, plants sprout from the stumps, root crowns, rhizomes, and/or aerial stems [294,296]. Salmonberry sprouting ability may decline after repeated, successive (i.e., monthly) top-kill [349], or when top-kill occurs between May and June [348]. Simulated response of canopy cover and height to manual cutting was compared at each phenological stage: establishment (early spring bud break in May), reproductive (early summer fruit set in late June and early July), and senescence (fall, October). The longest-duration reduction of cover occurred following cutting at the reproductive stage [198].

Approaches to control salmonberry include various types and combinations of site preparation, mechanical, biological, and chemical control (table 12). This information is beyond the scope of this review. The reader is encouraged to review the publications of interest in table 12, noting that some of the information included may be outdated and may not represent current best management practices. In addition, the reader is encouraged to consider all potential impacts of control methods (e.g., see Clarren 2014).

Table 12—Publications with information on salmonberry control.
Location Control method(s) Title Source
North America Chemical The role of herbicides for enhancing forest productivity and conserving land for biodiversity in North America Wagner et al. (2004) [334]
Pacific Northwest Chemical The effect of hexazinone, sulfometuron, metsulfuron, and atrazine on the germination success of selected Ceonothus and Rubus species Rose and Ketchum (2002) [262]
British Columbia Biological (fungi) Occurrence of Phoma argillacea on Rubus spectabilis in British Columbia and an evaluation of its potential as a forest weed biological control agent Sumampong et al. (2008) [300]
British Columbia Mechanical, site preparation Red alder-salmonberry complex: Operational summary for vegetation management British Columbia Ministry of Forests (2002) [28]
British Columbia Biological (fungi) Colonisation of leaves and twigs of Rubus parviflorus and R. spectabilis by endophytic fungi in a reforestation site in British Columbia Shamoun and Sieber (2000) [276]
British Columbia Biological (fungi) The development and potential role of mycoherbicides for forestry Wall et al. (1992) [335]
British Columbia Chemical Site preparation: chemical Otchere-Boateng and Herring (1990) [242]
British Columbia Chemical Effects of forest herbicides on some important wildlife forage species Balfour (1989) [17]
Washington, Oregon Chemical Interspecific competition and herbicide injury influence 10-year responses of coastal Douglas-fir and associated vegetation to release treatments Harrington et al. (1995) [118]
Oregon Site preparation Site preparation on coastal sites Stein (1990) [290]
Oregon Biological (domestic sheep) Seasonal diets of herded sheep grazing Douglas-fir plantations Leininger and Sharrow (1987) [8398]
Oregon Site preparation Brush control alternatives for forest site preparation Newton and Roberts (1977) [232]

APPENDIX


Table A1—Ecosystems, Associations, Cover Types, and BLM Regions where salmonberry occurs.
Ecosystems [87]
FRES20 Douglas-fir
FRES23 Fir–spruce
FRES24 Hemlock–Sitka spruce
FRES25 Larch
FRES27 Redwood
FRES28 Western hardwoods
FRES34 Chaparral-mountain shrub
Kuchler Plant Associations [178,179]
K001 Spruce–cedar–hemlock forest
K002 Cedar–hemlock–Douglas-fir forest
K003 Silver fir–Douglas-fir forest
K006 Redwood forests
K012 Douglas-fir forest
K014 Grand fir–Douglas-fir forest
K029 California mixed evergreen forest
K035 Coastal sagebrush
SAF Forest Cover Types [285]
210 Interior Douglas-fir
211 White fir
213 Grand fir
221 Red alder [216]
222 Black cottonwood–willow [59]
223 Sitka spruce [122]
224 Western hemlock
225 Western hemlock–Sitka spruce [218]
226 Coastal true fir–hemlock
227 Western redcedar–western hemlock [123]
228 Western redcedar [124]
230 Douglas-fir–western hemlock [255]
232 Redwood [263]
234 Douglas-fir–tanoak–Pacific madrone
SRM Rangeland Cover Types [163,183]
203 Riparian woodland
204 North coast shrub
901 Alder

Table A2—Common and scientific names of plants often associated with salmonberry. Follow the links to other FEIS Species Reviews. Nonnative plant species are indicated with an asterisk*.
Common name Scientific name
Ferns and fern allies
deer fern Blechnum spicant
feather mosses Hylocomium spp.
lady fern Athyrium filix-femina
western swordfern Polystichum munitum
western brackenfern Pteridium aquilinum
Forbs
American skunkcabbage Lysichiton americanus
Bohemian knotweed* Polygonum × bohemicum
coastal miterwort Mitella ovalis
false lily of the valley Maianthemum dilatatum
giant knotweed* Polygonum sachalinense
knotweed* Polygonum spp.
lupine Lupinus spp.
northern bedstraw Galium triflorum
Pacific golden saxifrage Chrysosplenium glechomifolium
Pacific waterleaf Hydrophyllum tenuipes
redwood-sorrel Oxalis oregana
seacoast angelica Angelica lucida
Siberian springbeauty Claytonia sibirica var. sibirica
tansy ragwort Senecio jacobaea
threeleaf foamflower Tiarella trifoliata
wall-lettuce* Mycelis muralis
Graminoids
bluejoint Calamagrostis canadensis
common velvetgrass* Holcus lanatus
reed canarygrass* Phalaris arundinacea
sedges Carex spp.
tufted hairgrass Deschampsia cespitosa
Shrubs
Alaska blueberry Vaccinium alaskaense
arctic raspberry Rubus arcticus
baccharis Baccharis spp.
blue elderberry Sambucus nigra subsp. cerulea
blueberry Vaccinium spp.
bunchberry dogwood Cornus canadensis
California blackberry Rubus ursinus
California hazelnut Corylus cornuta subsp. californica
California huckleberry Vaccinium ovatum
Cascara buckthorn Frangula purshiana
common snowberry Symphoricarpos albus
currant Ribes spp.
devilsclub Oplopanax horridus
elderberry Sambucus spp.
grayleaf raspberry Rubus idaeus subsp. strigosus
Himalayan blackberry* Rubus armeniacus
hollyleaved barberry Mahonia aquifolium
huckleberry Vaccinium spp.
mountain heather Cassiope spp.
northern black currant Ribes hudsonianum
oval-leaf huckleberry Vaccinium ovalifolium
Pacific poison-oak Toxicodendron diversilobum
red elderberry Sambucus racemosa
red huckleberry Vaccinium parvifolium
rose spirea Spiraea douglasii
salal Gaultheria shallon
Scotch broom* Cytisus scoparius
Sitka alder Alnus viridis subsp. sinuata
stink currant Ribes bracteosum
thimbleberry Rubus parviflorus
trailing black currant Ribes laxiflorum
vine maple Acer circinatum
willows Salix spp.
Trees
bigleaf maple Acer macrophyllum
bitter cherry Prunus emarginata
black cottonwood Populus trichocarpa
Douglas-fir
  coast Douglas-fir
  Rocky mountain Douglas-fir
Pseudotsuga menziesii
  Pseudotsuga menziesii var. menziesii
  Pseudotsuga menziesii var. glauca
Engelmann spruce Picea engelmannii
grand fir Abies grandis
mountain hemlock Tsuga mertensiana
Oregon ash Fraxinus latifolia
Pacific silver fir Abies amabilis
Port-Orford-cedar Chamaecyparis lawsoniana
red alder Alnus rubra
redwood Sequoia sempervirens
Sitka spruce Picea sitchensis
western hemlock Tsuga heterophylla
western larch Larix occidentalis
western redcedar Thuja plicata

Table A3—Vegetation classifications and related publications that include salmonberry as an important, indicator, or dominant species.
Location Title Citation
Pacific Northwest
Oregon and Washington Natural vegetation of Oregon and Washington Franklin and Dyrness (1988) [81] (1973) [80]
western Oregon, southwestern Washington Major indicator shrubs and herbs on national forests of western Oregon and southwestern Washington Halverson (1986) [102]
throughout Ecoclass coding system for the Pacific Northwest plant associations Hall (1984) [100]
California
northern coast Status of vegetation classification in redwood ecosystems Mahony and Stuart (2007) [194]
Oregon
northwestern Oregon Field guide to riparian plant communities in northwestern Oregon McCain and Christy (2005) [201]
Oregon Regional gradient analysis and spatial pattern of woody plant communities of Oregon forests Ohmann and Spies (1998) [237]
southwestern Oregon Field guide to the forested plant associations of southwestern Oregon Atzet et al. (1996) [12]
central Coast Ranges The structure and dynamics of red alder communities in the central Coast Range of western Oregon Carlton (1988) [44]
Valley of the Giants, northwestern Oregon A guide to the vegetative communities at the Valley of the Giants, Outstanding Natural Area, northwestern Oregon, USA Sonnenfeld (1987) [286]
Siuslaw National Forest Plant association and management guide: Siuslaw National Forest Hemstrom and Logan (1986) [130]
western Cascade Range Riparian vegetation in Oregon's western Cascade Mountains: composition, biomass, and autumn phenology Campbell and Franklin (1979) [42]
western Oregon Plant succession in the Alnus rubra/Rubus spectabilis habitat type in western Oregon Henderson (1978) [131]
Oregon coast Synecological features of a natural headland prairie on the Oregon coast Davidson (1967) [57]
Washington
eastern Washington Classification and management of aquatic, riparian, and wetland sites on the national forests of eastern Washington: series description Kovalchik and Clausnitzer (2004) [176]
Olympic National Forest Indicator species of the Olympic National Forest Lesher and Henderson (1989) [189]
Mount Rainier National Park The forest communities of Mount Rainier National Park Franklin et al. (1988) [82]
North Cascades National Park Forest types of the North Cascades National Park Service Complex Agee and Kertis (1987) [3]
Olympic National Park A taxonomical - ecological study of the vegetation by habitats in eight forest types of the Olympic Rain Forest, Olympic National Park, WA Sharpe (1956) [277]
Alaska
Kenai Fjords National Park Landcover classes, ecosystems and plant associations: Kenai Fjords National Park Boggs et al. (2008) [22]
Tongass National Forest, Petersburg and Wrangell Districts Constancy and cover of plants in the Petersburg and Wrangell Districts, Tongass National Forest and associated private and other public lands, southeast Alaska Mead (2002) [206]
Copper River Delta Classification of community types, successional sequences, and landscapes of the Copper River Delta, Alaska Boggs (2000) [21]
Chugach National Forest Plant community types of the Chugach National Forest: Southcentral Alaska DeVelice et al. (1999) [66]
Tongass National Forest, Chatham Area Forest plant association management guide: Chatham Area, Tongass National Forest Martin et al. (1995) [195]
Tongass National Forest, Ketchikan Area Forest plant association management guide: Ketchikan Area, Tongass National Forest DeMeo et al. (1992) [64]
southeastern Alaska Vegetation and environment in old growth forests of northern southeast, Alaska: A plant association classification. Martin (1989) [196]
western Alaska Characteristics of some grassland, marsh, and other plant communities in western Alaska Hanson (1951) [111]
British Columbia
throughout Indicator plant species in Canadian forests Ringius and Sims (1997) [258]
throughout Classification of natural forest communities of coastal British Columbia, Canada Klinka et al. (1996) [172]
throughout Indicator plants of coastal British Columbia Klinka et al. (1989) [169]

Table A4—Common and scientific names of wildlife species mentioned in this review. Follow the links to FEIS Species Reviews.
Common name Scientific name
Birds
American robin Turdus migratorius
band-tailed pigeon Patagioenas fasciata
Cassin's auklets Ptychoramphus aleuticus
gray catbird Dumetella carolinensis
rufous hummingbird Selasphorus rufus
varied thrush Ixoreus naevius
Mammals
American beaver Castor canadensis
American black bear Ursus americanus
Columbian white-tailed deer Odocoileus virginianus leucurus
coyote Canis latrans
elk Cervus elaphus
grizzly bear Ursus arctos horribilis
moose Alces americanus
mountain beaver Aplodontia rufa
mountain goat Oreamnos americanus
mule deer
  Columbian black-tailed deer
  Sitka black-tailed deer
Odocoileus hemionus
  Odocoileus hemionus columbianus
  Odocoileus hemionus sitkensis
North American porcupine Erethizon dorsatum
Roosevelt elk Cervus elaphus roosevelti
white-footed vole Arborimus albipes

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