Index of Species Information

SPECIES:  Gaultheria shallon

Introductory

SPECIES: Gaultheria shallon
AUTHORSHIP AND CITATION : Tirmenstein, D. 1990. Gaultheria shallon. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].
ABBREVIATION : GAUSHA SYNONYMS : NO-ENTRY SCS PLANT CODE : GASH COMMON NAMES : salal Oregon wintergreen TAXONOMY : The currently accepted scientific name of salal is Gaultheria shallon Pursh (Ericaceae) [72]. LIFE FORM : Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY


DISTRIBUTION AND OCCURRENCE

SPECIES: Gaultheria shallon
GENERAL DISTRIBUTION : Salal grows along the Pacific Coast inland to the western slope of the Cascades and Coast Ranges [67].  It occurs from southeastern Alaska and central British Columbia southward to southern California [28,67,102,130]. ECOSYSTEMS :    FRES20  Douglas-fir    FRES24  Hemlock - Sitka spruce    FRES27  Redwood    FRES28  Western hardwoods STATES :      AK  CA  OR  WA  BC BLM PHYSIOGRAPHIC REGIONS :    1  Northern Pacific Border    2  Cascade Mountains    3  Southern Pacific Border KUCHLER PLANT ASSOCIATIONS :    K001  Spruce - cedar - hemlock forest    K002  Cedar - hemlock - Douglas-fir forest    K006  Redwood forest    K029  California mixed evergreen SAF COVER TYPES :    221  Red alder    224  Western hemlock    225  Western hemlock - Sitka spruce    226  Coastal true fir - hemlock    227  Western redcedar - western hemlock    228  Western redcedar    229  Pacific Douglas-fir    230  Douglas-fir - western hemlock    231  Port Orford cedar    232  Redwood    234  Douglas-fir - tanoak - Pacific madrone SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Salal grows as an understory dominant in a variety of lowland to montane, coniferous or mixed evergreen forests.  Common overstory dominants include Douglas-fir (Pseudotsuga menziesii), western hemlock (Tsuga heterophylla), Sitka spruce (Picea sitchensis), lodgepole pine (Pinus contorta), western redcedar (Thuja plicata), tanoak (Lithocarpus densiflorus), and Pacific silver fir (Abies amabilis).  Evergreen huckleberry (Vaccinium ovatum), red huckleberry (V. parvifolium), Sadler oak (Quercus sadleriana), rhododendron (Rhododendron spp.), vine maple (Acer circinatum), oceanspray (Holodiscus discolor), bracken fern (Pteridium aquilinum), dwarf Oregon grape (Mahonia nervosa), salmonberry (Rubus spectabilis), California hazel (Corylus cornuta), western swordfern (Polystichum munitum), deerfern (Blechnum spicant), threeleaf foamflower (Tiarella unifoliata) are common understory codominants. Salal is listed as a dominant or indicator in the following publications: Forest types of the North Cascades National Park Service Complex [1] Description and classification of the forests of the upper Illinois   River drainage of southwestern Oregon [4] Forest associations and secondary succession in the southern Oregon   coast Range [8] Plant communities and environmental interrelationships in a portion of   the Tillamook Burn, northwestern Oregon [9]      Classification of montane forest community types in the Cedar River   drainage of western Washington, U.S.A. [26] Vegetation of the Douglas-fir region [34] Plant association and management guide for the western hemlock zone: Mt.   Hood National Forest [51] Plant association and management guide for the Pacific silver fir zone,   Mt. Hood and Willamette National Forests [60] Plant association and management guide: Willamette National Forest [62] Indicator plants of British Columbia [79] Biogeoclimatic ecosystem classification of British Columbia [108]

MANAGEMENT CONSIDERATIONS

SPECIES: Gaultheria shallon
IMPORTANCE TO LIVESTOCK AND WILDLIFE : Browse:  In many areas, salal is browsed at least moderately by deer and elk [28,53,113].  However, use varies geographically as well as seasonally.  Salal is heavily browsed by black-tailed deer on the Queen Charlotte Islands of British Columbia [45].  Persistent leaves enhance winter value, and in many areas, including the Oregon Coast Range, salal is an important winter food for black-tailed deer and mule deer [14,15,65,100,103].  Deer use is often heaviest when other low-growing species become covered with snow [64,65].  High elevation stands are generally not used by deer in winter [64].  Seasonal black-tailed deer use has been documented as follows in western Washington [14]:                   percent total volume Jan.  Feb.   March  April  May  June  July  Aug.  Sept.  Oct.  Nov.   Dec. 30.4  12.3   15.2   12.9   1.1  0.5   3.9   17.2  1.0    5.1   --     27.6 Roosevelt elk consume some salal browse, particularly during the winter months [8,53].  Light to moderate elk use has also been reported during fall and spring in some areas [116], but elsewhere, browse may be ignored during spring and summer [53].  Winter elk use may occasionally be locally heavy [116].  Salal is considered an important "emergency" food in some locations [128]. Small mammals such as the mountain beaver also feed on salal [128]. This shrub is a preferred food of the mountain beaver in parts of the western Cascades [68].  Leaves make up a small portion of the white-footed vole's July diet in parts of Oregon [133]. In some areas, domestic sheep and goats browse salal [113]. Fruits and flowers:  Salal fruit is readily eaten by many birds and mammals [67].  The band-tailed pigeon, wrentit, ruffed, spruce, and blue grouse, and numerous songbirds feed on "berries" when available [28,87,138].  In some areas, blue grouse chicks exhibit a marked preference for salal fruit, and both chicks and adults consume large numbers during July and August [77].  Some hummingbird use of flowers has also been reported [107].  Black-tailed deer of western Washington consume the flowers of salal [14].  Mammals such as the red squirrel, black bear, black-tailed deer, Townsend's chipmunk, and Douglas' squirrel also feed on salal fruit [45,87]. PALATABILITY : Salal browse is at least moderately palatable to many big game species, but relatively unpalatable to domestic livestock [128].  In some locations, leaves are readily eaten by black-tailed deer [23].  Deer often exhibit a marked preference for tender sprouts on burned-over sites [113].  Evergreen foliage remains palatable during the winter months.  Overall palatability of salal has been rated as follows [64,113,116]:                         CA            OR             WA Cattle            poor to useless     ----          ---- Domestic sheep    poor                ----          ---- Horses            useless             ----          ---- Elk               ----                ----          fair Deer              fair to poor        moderate      ---- Domestic goats    fair to poor        ----          ---- Salal fruit is palatable to a wide variety of birds and mammals. NUTRITIONAL VALUE : Browse:  The nutrient content of salal browse varies according to plant part and with the stage of phenological development.  However, in general browse has relatively low nutritional value.  Black-tailed deer which fed exclusively on salal browse exhibited signs of malnutrition [64].  Nutrient content has been documented as follows [14,111]: crude       ether       crude       N-free      total       Ca protein     extract     fiber       extract     ash                               (percent)   6.75         5.19       21.78        58.23      6.65        1.203 Mg          K           PO4 0.434       0.572       0.272                   average percent weight -                   N      P       Mg    Ca     Na         K              stem             .25     .05     .05   .18    .0010     .24 foliage          .81     .08     .21   .81    .0030     .40 Fruit:  Nutrient value of salal fruit is listed below [101]:             kjoules     calories    protein   carbo.    ash     lipid               x 1,000                 (g)         (g)     (g)     (g) fresh       15.52        3.71       0.13      0.79      0.03     0.05 dried       14.69        3.51       0.06      0.88      0.04     0.01             Ca         Fe         Mg             (mg)       (mg)       (mg) fresh       3.77       0.04       0.91 dried       3.44       0.04       0.21 COVER VALUE : Salal provides important cover for a variety of wildlife species [27]. Western hemlock/dwarf Oregon grape-salal, western hemlock/vine maple-salal, and Sitka spruce-salal communities offer good hiding cover for deer and elk, although dense shrub development can sometime limit big game use [61,127].  Red huckleberry-salal shrubfields protect black-tailed deer from winter winds [65]. VALUE FOR REHABILITATION OF DISTURBED SITES : Once established, salal spreads aggressively and is well-suited for use as a ground cover on erosive banks, roadcuts, highway right-of-ways, and other types of reclaimed ground [80,129].  It can also aid in stabilizing coastal dunes and in protecting vulnerable watersheds [28]. Salal may be propagated by seed [28,80].  Cleaned seed averages 3,209,000 per pound (7,068/kg) and remains viable for "moderate periods" when properly stored [28].  Seed is generally sown in winter or spring [138].  Seedlings exhibit slow growth, but propagation from seed is generally the most economical means of growing salal [129].  Seed collection, handling, and planting methods have been considered in detail [28,80,129]. Salal can also be propagated vegetatively from root, stem, or rhizome cuttings, although propagation can be difficult and initial growth slow [28,129].  Best results are generally obtained from cuttings taken in late summer [80].  Salal can also be propagated by layering, or from suckers and stolons [28].  Various modes of vegetative propagation have been examined in detail [28,80,129]. OTHER USES AND VALUES : Fruit of salal was traditionally utilized by many native peoples of the Northwest [101].  The spicy fruit was eaten fresh, dried, or mashed into cakes [28,50,130].  Leaves were dried, mixed with kinnikinnick (Arctostaphylos spp.) and smoked [28,50].  Teas made from the leaves were used to treat coughs, tuberculosis, and diarrhea [50]. Salal is cultivated as an ornamental.  Plants are used in landscaping [50] and serve as an excellent ground cover [66].  Salal can be used to attract wildlife species to backyard gardens [80].  The attractive foliage is used by florists under the name "lemon leaf" as an addition to cut flowers [28,87,113]. The sweet, "bland but pleasant" fruit can be used alone or mixed with other wild berries to make jellies or preserves [28,80].  Approximately 8 minutes of harvesting is required to collect 0.44 pint (250 ml) of fruit [83].  Many species of Gaultheria contain oil of wintergreen and can be used as flavoring agents [113]. OTHER MANAGEMENT CONSIDERATIONS : Timber harvest:  Salal commonly increases after timber harvest [2,61,89].  Generally, if present in the understory prior to harvest, it will also form part of the postdisturbance community [22].  Heavy thinning can increase salal biomass by up to 2.8 times [117].  The effects of timber harvest on salal have been examined in a number of studies [8,30,31,57,70,71,73,79,94,112,115,131]. Competition:  Salal competes vigorously with conifer regeneration in some locations [79].  On moist sites, this shrub commonly competes with Douglas-fir, Sitka spruce, and western hemlock, and to a lesser degree with western redcedar [143].  In general, the nutrient-demanding Sitka spruce is most harmed by competition with salal [89], but salal can also significantly reduce the basal area and stocking of Douglas-fir seedlings on some sites [16].  In some areas, salal vigorously competes with Douglas-fir for both water and nutrients [15,41,104] resulting in poor seedling growth [134].  In many problem areas, soil moisture deficits are common during the growing season, and competition for moisture may be of primary significance [109].  Competition is often pronounced in drier low elevation forests of coastal British Columbia where dense thickets of salal commonly form on cutover sites [24,42]. Growth of forest crop trees is commonly reduced at approximately 6 to 8 years after planting in coastal Sitka spruce-western hemlock-western redcedar, and western hemlock forests where a dense ground cover of salal is present [92,134].  This growth check period may be due to the direct effects of competition with salal or allelopathy associated with this ericaceous shrub [134].  Anderson [3] reports that a dense growth of salal can also inhibit regeneration of maples (Acer spp.), as samaras are physically prevented from reaching the forest floor [3]. On some sites in western Washington, salal may actually add nutrients to the soil and apparently has no adverse effect on the growth of Douglas-fir [45,79].  Klinka and others [79] report that the amount of nitrogen tied up by salal is relatively small and is not likely to be critical for tree growth except on very poor sites.  In some areas, conifer regeneration is typically better on sites dominated by salal than on sites dominated by western swordfern or vine maple [29]. Still, much research has focused on ways to eliminate salal to improve conifer regeneration.  Recommendations for minimizing salal competition with conifer seedlings include [89]:            (1) preventing fires on naturally regenerated clearcuts            (2) preparing seedbeds to encourage prompt natural                regeneration            (3) planting seedlings immediately after timber removal;                           adding fertilizers where necessary Successive light treatments may be preferable to a single heavy tree removal [104].  When thinning, particular care should be taken to avoid creating large gaps in the canopy [104].  It may be desirable to maintain greater stand density on dry sites with salal present [104]. Competition between conifer seedling and salal occurs largely below ground [16], and seedlings should be planted as early as possible after timber harvest to allow seedlings a "head start" [16].  In some areas, planting densities necessary to shade out salal quickly are impractically high [16].  Models have been developed which explore the effects of salal competition on the growth of various conifer seedlings [88].  The effects of competition have been considered in detail [16,41,88,89,92,104,135,136,142,143].  However, in many instances, elimination of salal is difficult, uneconomical, or impractical. Bunnell [15] reports that "...attempts to reduce salal abundance may be unwarranted; the species appears well adapted to persist." Chemical control:  Salal is resistant to many herbicides including 2,4-D, velpar, 2,4,5-T, amitrole, picloram, and silvex [12,106,120]. Site characteristics [24,41] and season and mode of application can greatly influence the response of salal to herbicides [121].  Repeated application of Garlon is effective although often impractical [24] or prohibitively expensive.  Silvex can also be relatively effective in reducing cover when properly applied [121].  Salal appears to be most susceptible to foliage sprays in diesel oil carriers when applied at budbreak [121].  Plants are less seriously damaged by herbicides applied late in the growing season or by those applied in water or oil-in-water emulsions [121].  In test applications, few of the damaged salal plants were actually killed by herbicides, and recovery was generally rapid [121].  However, herbicides can sometimes produce sufficient control for conifer release [121].  Detailed information on the response of salal to herbicides is available [12,17,21,24,45,120,121]. Mechanical removal:  Various types of mechanical removal or soil disturbance can stimulate sprouting of salal and produce increased cover [45].  As rhizomes are broken, new plants commonly form [15].  Harvest techniques which disrupt rhizomes, such as the use of skidders, can produce additional management problems by fostering the spread of salal [15].  In coastal British Columbia, spot scarification appears to be relatively ineffective in producing long-term control of salal [24]. Pretreatment levels can be reached by the third growing season [24]. Blade scarification was more effective, reducing cover to 6 percent but resulted in significant site degradation [24].  Details on mechanical treatments are available [24,41,135,136]. Biomass:  In general, aboveground biomass of salal appears to be inversely proportional to the amount of overstory foliage [85]:                               stand age                               (years)                       22          30          42          73 salal biomass (kg/ha)   6300.6      4112.2      3394.0      1010.2 Heavy fertilizer application can decrease the aboveground biomass of salal [117]. Wildlife:  Salal fruit production may be limited beneath a closed canopy [15].  Disturbances which eliminate portions of the overstory presumably increase fruit production.  Where management goals are aimed at increasing winter big game forage, evidence suggests that salal will respond favorably to thinning [15]. Research indicates that mountain lion, coyote, and wolf urine can be used to inhibit or stop deer use of salal browse [124]. Livestock:  Salal is susceptible to trampling damage [102]. Chemical composition:  Evidence suggests that salal may be somewhat allelopathic [25,136,141].  The foliage and roots of salal are resistant to decay and can reduce decomposition and water availability [79].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Gaultheria shallon
GENERAL BOTANICAL CHARACTERISTICS : Salal is an erect to spreading, clonal evergreen shrub or subshrub which grows 1.3 to 10 feet (0.4-3 m) in height [16,67,80,98].  This loosely to densely branched shrub often forms dense, nearly impenetrable thickets [45].  Stems are pilose to hirsute [67] and branchlets glandular to pubescent [98].  Twigs are reddish-brown with shredding bark [130]. Most biomass is concentrated below ground [102] and an extensive, but variable network of roots and rhizomes [24] occupies the top layer of soil [79]. Leaves are ovate to ovate-elliptic, sharply serrulate, and 2 to 4 inches (5-10 cm) in length [66,80].  The shiny dark green, alternate leaves are thick and leathery [24,45,80]. Small, urn-shaped flowers are borne in showy clusters on terminal and subterminal bracteate racemes [42,45,66,80].  The white, pink or deep-rose tinged flowers are sticky and glandular [80,98].  Floral morphology has been examined in detail [19].  Fruit is a round, reddish, purplish, or bluish black "pseudoberry" or capsule which is made up of a fleshy outer calyx [45].  Fruits are covered with tiny hairs [42] and average 0.24 to 0.4 inch (6-10 mm) in diameter [66].  Each fruit contains an average of 126 brown, reticulate seeds approximately 0.04 inch (1 mm) in length [45,98]. Salal leaves generally live for 2 to 4 years [45].  Twigs survive for 16 years or longer, but bear leaves only during the first few years [45]. Rhizomatous portions of individual plants can live for hundreds of years [102]. RAUNKIAER LIFE FORM :       Phanerophyte REGENERATION PROCESSES : Salal is capable of reproduction from seed and vegetative regeneration. However, seedling establishment is apparently insignificant where plants are already established.  Additional expansion of existing clones occurs through layering, sprouting of rhizomes, root suckering, and sprouting from the stem base [45]. Seed:  Good seed crops are produced regularly, except under a dense forest canopy where little or no seed is produced [45].  In a British Columbia study, only 8.7 percent of all twigs produced flowers, and no flowers were noted where the canopy cover exceeded 33 percent. Flowering beneath a forest canopy was limited to shoots more than 4 years of age.  Plants that flowered were, on the average, larger and more vigorous than those that did not.  Flowering characteristics were documented as follows [15]:                   age of shoot (years)                         < or = 4       5        6        7        8     flowering/total       shoots                0/13      5/37     8/53     2/7       3/10       new twigs             0/15     20/139   13/200   14/81     16/288 length of flowering       twig (cm)             ----      9.1     7.3      6.6       5.5 # of flowers per twig       ----      6.6     6.6      5.9       5.5                   Salal flowers are pollinated by insects such as bees and flies [45]. Seeds are dispersed by a variety of birds and mammals [45,118]. Evidence suggests that seeds consumed by bears may germinate more readily than uneaten seeds [102]. Germination:  Germination of salal is generally good under laboratory conditions, with up to 73 percent of the seed eventually germinating [28,80].  In other laboratory tests, average germination of 27 to 35 percent has been reported [45].  Stratification is not essential for germination [95], but periods of light (at least 8 hours per day), are [28].  In laboratory tests, seeds typically begin germinating within 27 [95] or 30 to 45 days [129].  Viability in storage appears limited [106].  Germination capacity declined from 31 to 21 percent after 1 year in storage at 40 degrees F (4 degrees C) but averaged 73 and 27 percent after 3 years in storage at 40 degrees F (4 degrees C) and room temperature, respectively [28]. Seedling establishment:  Potential for reproduction from seed appears poor under natural conditions [47,102].  Few seedlings establish despite the large numbers that germinate.  Seedling establishment may be limited to favorable microsites or to periods of unusual weather conditions [102].  Initial seedling growth is slow [45].  Seedlings may require 2 to 3 years to reach 3 to 5 inches (8-13 cm) in height [80].  Early seedling growth is favored by moist, acidic conditions and partial shade [28]. Seed banking:  Seed remains viable for several years when properly stored, but viability is probably much lower under natural conditions [45].  Kellman [74] sampled soil and litter from beneath 100-year old Douglas-fir-western hemlock stands in coastal British Columbia.  Core samples were divided into an upper layer, 0 to 2 inches (0-5 cm), and lower layer, 2 to 4 inches (5-10 cm).  Although seed was found in only 1 out of 34 cores, subsequent establishment did occur in laboratory tests [74].  Seed banking, although possible, is presumably a relatively unimportant regenerative strategy in salal. Vegetative regeneration:  Salal sprouts prolifically from roots, rhizomes, underground stems, and the stem base after disturbances which damage or remove aboveground plant parts [80,102,106,121], and expands through spreading roots and rhizomes in the absence of disturbance [24,102,106].  Layering, rooting at the stem nodes, and spread through stolons has also been reported [24,28].  Stems which are forced into the organic mat typically generate adventitious roots.  Salal plants are often made up of several individual aboveground shoots connected belowground by several meters of rhizomes [15]. Vegetative regeneration occurs under either a sparse or dense overstory canopy, and where canopy cover exceeds 33 percent, represents the only mode of regeneration.  Plants growing beneath a sparse overstory produced an average of 0.21 shoots per plant per year while those beneath a closed canopy generated 11 new shoots per plant per year. However, shoots typically live longer (10.33 years) beneath a sparse overstory canopy than beneath a closed canopy (6.25 years).  As the overstory canopy becomes more dense, investment in rhizome extension increases.  This expansion could represent an "escape from shading" under conditions of changing canopy gaps.  Bunnell notes that "under canopy, the spatial pattern of...shoots was better adapted to maintain plant persistence than to colonize new areas" [15].  Messier and others report that plants allocate greater energy to the rhizomes as they mature [92].  Bunnell observed that vegetative regeneration typically declines with increasing age (> 3 years) [15].  No new shoots were produced by plants 9 years or older.  Early sprout growth may be slow. Plants may need as long as 5 years to regenerate stems and produce aboveground growth [80]. SITE CHARACTERISTICS : Salal grows in warm, moist to dry, montane to lowland coastal conifer forests of the Pacific Northwest [50,66].  It occurs in a variety of communities including marginal peatland forests, soligenous fens, forested swamps, bogs, and muskegs [39,99,125].  In parts of British Columbia, it occurs in shrub communities at the driest edges of bogs [59,132].  Salal is tolerant of salt spray and commonly forms dense stands in northern coastal shrub communities [50,58].  It grows well on stabilized dunes, exposed slopes, rocky bluffs, and knolls near the ocean [37,45].  It is a common component of swampy shore pine or spruce woodlands [37,126].  Salal commonly grows vigorously after stands are opened by timber harvest and persists in many coastal brushfields. Salal grows well in partial shade, although vigor may be poor beneath a dense canopy [45].  This shrub persists in sun or shade [138].  Salal commonly forms dense thickets beneath the forest canopy and at forest margins [80].  In pygmy forests dominated by bishop pine (Pinus muricata), lodgepole pine, and cypress (Cupressus pygmaea), it grows as a dwarf, spreading shrub [126,137]. Salal typically occurs on moderately warm dry sites in western hemlock communities [26,51,79] and on very dry to wet sites in coastal Douglas-fir communities [100].  Salal grows on warm, dry sites with Pacific silver fir [60] and on drier sites in Port-Orford-cedar and tanoak communities [5,6].  It grows as an understory dominant in coastal coniferous forests [45] commonly dominated by western hemlock, western redcedar, Port-Orford cedar, Sitka spruce, lodgepole pine, and Alaska cedar (Chamaecyparis nootkatensis) [39,46,130,138].  Salal is also common in mixed evergreen, redwood (Sequoia sempervirens), and subboreal spruce communities, and in pygmy forests of northern California [52,98,114,126,137]. Plant associates:  Salal commonly occurs with species such as red alder (Alnus rubra), salmonberry (Rubus spectabilis), vine maple, western swordfern, rhododendron, vaccinium (Vaccinium spp.), dwarf Oregon grape, Pacific dogwood (Cornus nuttallii), tanoak, threeleaf foamflower, and deerfern in western hemlock or western hemlock-western redcedar forests [8,51,56,63].  Vine maple, oceanspray, dwarf Oregon grape, Pacific rhododendron (Rhododendron macrophyllum), and California hazel are common associates in Douglas-fir forests [3,44,48].  The understory may be depauperate in old growth stands.  In redwood forests, salal grows with dwarf Oregon grape, evergreen huckleberry, willow (Salix spp.), California hazel, Pacific madrone (Arbutus menziesii), California laurel (Umbellularia californica), and rhododendron [113,126].  In northern coastal scrub, chaparral broom (Baccharis pilularis), many-colored lupine (Lupinus varicolor), trailing blackberry, pearly everlasting, common velvetgrass (Holcus lanatus), and California oatgrass (Danthonia californica) are common associates [58]. Soil:  Salal grows on a variety of mineral and organic substrates including shallow rocky soils, sand dunes, coarse alluvium, glacial till, and peat [45,56].  Growth is generally best on moist sandy or peaty soils where salal occurs as a vigorous upright shrub [45].  Salal grows on nutrient poor to moderately rich soils [45,79].  On shallow, droughty soils, plants may assume a matlike growth form.  Salal commonly grows on decaying wood and stumps and can grow as an epiphyte on living trees in extremely humid areas [45].  It occurs on soils derived from a wide range of parent material including diorite, breccia and basalt, serpentine, granite, and metamorphic rock [51,114,138,139]. Climate:  This shrub grows in hypermaritime to maritime zones characterized by cool, humid to perhumid, mesothermal climate [78,79]. Winters are typically mild with little snow accumulation [45].  Plants are dwarfed in drier areas [138].  Salal reaches greatest size and abundance in the fogbelt along the Pacific Coast [129].  Plants are sensitive to frost [45]. Elevation:  Salal typically grows at low to intermediate elevations. Elevation by geographic location is as follows [45,98]:                   > 2,500 feet (> 763 m) in CA                   0 to 2,624 feet (0-800 m) in s coastal BC                   < 33 to 116 feet (<100-200 m) in n coastal BC SUCCESSIONAL STATUS : Salal is a residual species which persists on many types of newly disturbed sites [31,36,47,71].  It can rapidly colonize open areas, particularly on undisturbed soil [15,31] and appears well adapted for "opportunistic survival in ...changing canopy gaps" [15].  Salal commonly increases in abundance and cover on clearcuts in old growth western hemlock and western hemlock-western redcedar forests of the Northwest [136].  Typically, it is initially much reduced by logging and postharvest fires but recovers dramatically [47].  Salal is a common constituent of persistent seral brushfields and can remain dominant for 25 years [36] or more.  The shrub was observed on mudflow channels, buried roadbanks, blowdown, and scorch sites soon after the eruption of Mount St. Helens [49,91]. Douglas-fir-western hemlock: Salal grows in early seral to climax stands in Douglas-fir-western hemlock forests and in coastal western hemlock forests of the Northwest [42,55,56,78].  Weedy invaders such as groundsel (Senecio spp.), fireweed (Epilobium angustifolium), pearly everlasting (Anaphalis margaritacea), and bracken fern are common dominants during the first three growing seasons after fire or other disturbances [61,82,84].  Subsequent recovery of salal is commonly rapid [8] with this shrub assuming prominence within 3 to 5 years after disturbance [39,122].  According to Bunnell, 85 percent of the space that will be occupied by salal is occupied within 3 years [15].  By year 8, salal can fully occupy the belowground environment [143] and continues to increase as fireweed declines [8].  By the 10th growing season, salal may reach 2 to 3 feet in height [112].  In western Washington, salal commonly increases in density as second growth conifers begin to overtop the shrub layer [14].  Salal commonly shares dominance with dwarf Oregon grape during postdisturbance years 7 to 50 in the Oregon Coast Range [8]. Salal is a principal understory species in many Douglas-fir forests of the Olympic Mountains where it dominates 65- to 90-year-old and 300-year-old stands [34].  It is common in second growth Douglas-fir stands of the Oregon Coast Range [8] and northeastern Olympic Mountains of western Washington [105], but in some areas, it may be sporadic or absent in the shaded understory of immature, closed canopy stands [79]. Salal can attain temporary dominance approximately 22 years after disturbance [84].  Salal commonly attains peak abundance in middle-late to late seral stages following fire [47].  As the overstory develops further, cover gradually declines [84].  Cover of salal by stand age has been documented as follows in western Washington [84]:             stand age (years)             (percent ground cover) -       5      22     30      42      73    12.22   65.26   44.56   43.72   30.90 Cover was documented as follows in a Douglas-fir-western hemlock forest of western Cascades of Oregon [115]:                               years (percent cover)        2     5     10     15     20     30     40    undist. old growth      7.37   1.41   10   8.52   9.93   17.74  14.97         7.37 Salal commonly persists as an understory dominant or codominant in relatively dry Douglas-fir forests of British Columbia [79] and the Pacific Northwest [5].  However, many moist northwestern Douglas-fir forests are seral to western hemlock types, and with time, the composition of the overstory gradually shifts from Douglas-fir to hemlock or cedar-hemlock [39].  True climax status may not be reached for several hundred years [84].  In climax stands, the herbaceous layer is often depauperate [8].  Salal and dwarf Oregon grape are often the only two species with more than 1 percent cover [8].  Salal is a common understory dominant in climax western hemlock-western redcedar forests [8] and in coastal western hemlock forests [78] but may be absent in mature western redcedar forests of coastal British Columbia [79]. Port-Orford-cedar:  Salal occurs in seral to climax stands in Port-Orford cedar communities [5].  It occurs as an understory dominant in drier Port-Orford-cedar forests of the Siskiyou Mountains [6]. Redwood, Sitka spruce:  Salal commonly increases after logging in redwood [11] and Sitka spruce [2] forests. SEASONAL DEVELOPMENT : Salal exhibits variable annual and geographic phenological development. In a Washington study, bud burst occurred in April, with rapid vegetative growth occurring from April until early June when growth peaked [45]. Plants generally flower in late spring or early summer [80] with fruit ripening from August through October [28,45].  Fruit may persist on the stem until December [45,138].  Generalized flowering and fruiting dates by geographic location are as follows: location          flowering        fruiting          authority AK                May-June         ----              [130] BC                June 12-July 4   June-September    [45,83]  CA                March-July       ----              [113] w WA              ----             3rd week of June  [45] w OR, sw WA       May-July         ----              [50] Northwest         May-July         ----              [67]    

FIRE ECOLOGY

SPECIES: Gaultheria shallon
FIRE ECOLOGY OR ADAPTATIONS : The shade-tolerant salal appears well able to persist under a regime of relatively infrequent fires.  Long fire-free intervals are common in many climax coastal coniferous forests of the Pacific Northwest [60]. Fire occurs infrequently in most coastal western hemlock forests due to marine climatic influences [5].  Western hemlock-Douglas-fir forests codominated by salal and dwarf Oregon grape commonly burn at approximately 320-year intervals [105].  Fire intervals in tanoak-salal/dwarf Oregon grape communities of the western Siskiyous have been estimated at 60 years [5].  While inland redwood forests burn every 26 to 52 years, coastal redwood forests experience fires at 50 to 500-year intervals [123].  In western Oregon, Douglas-fir/oceanspray -salal communities are common on sites which have been lightly burned during the past 200 years.  Salal, because of its prolific sprouting ability, can also survive shorter fire-free intervals.  In western Oregon, bracken fern-salal communities commonly develop on frequently burned sites [8]. Salal generally sprouts from the roots, rhizomes, or stem base after aboveground vegetation is damaged or consumed by fire.  Birds and mammals may disperse some seed from off-site.  Limited reestablishment through seed may occur, although vegetative regeneration is apparently the dominant mode of reestablishment [47]. POSTFIRE REGENERATION STRATEGY :    Tall shrub, adventitious-bud root crown    Rhizomatous shrub, rhizome in soil

FIRE EFFECTS

SPECIES: Gaultheria shallon
IMMEDIATE FIRE EFFECT ON PLANT : Salal is described as a woody survivor [140].  Underground portions of the plant commonly survive even when aboveground vegetation is consumed by fire [16,57].  Portions of the stem base also survive many low severity fires [57].  Hot burns on dry, shallow soil can result in lethal heat penetration to underground regenerative structures [45]. "Moderate damage" has been reported after light burns [5]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Vegetative response:  Salal typically sprouts readily from the roots, rhizomes, or stem base after light to moderate fires [5,16,79].  Fires of light to moderate intensity stimulate sprouting, but more intense fires can damage underground regenerative structure and reduce or eliminate sprouting [45]. Seed:  Postfire reestablishment through seed appears to be relatively unimportant in salal [47]. Postfire recovery:  Recovery of salal varies according to fire intensity and severity [48].  Rhizome expansion can be rapid [136] or relatively slow depending on the amount of damage received [57,140].  Plants are often observed soon after fire [68,76,82] but may only develop slightly during the first year [68].  Following a moderate burn in British Columbia, salal was present during the first growing season and increased in abundance by the third growing season [82].  However, few plants were observed during the first growing season after an intense fire in the same area [82].  By the 3rd year after this fire, only small, scattered colonies of salal were present [82].  Bailey [8] observed increases in cover by the 8th year after logging and fire in western Oregon.  Salal can become dominant within 10 years after fire in parts of British Columbia [102].  Salal can reach 2 to 3 feet (0.6-0.9 m) in height by the tenth growing season after fire [112].  Recovery was documented as follows after logging and fire in the Oregon Coast Range [122]:                   before burn             1 year after burn             orig.       seedlings         orig. stems       seedlings             stems                         + sprouts                              (# per acre) N-aspect     250             0                640                 0 S-aspect   2,840             0             15,960                 0 Following fire in British Columbia, cover reached 18 percent after 2 years and had increased to 55 percent with 8 years [141].  However, 4 years after intense summer wildfires in the North Cascades of Washington, cover of salal on two sites ranged from 0.7 to 1 percent [93]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : Timber harvest:  Evidence indicates that postfire recovery of salal on some harvested sites may be delayed by slash-burning [94,131].  Recovery may be particularly slow after hot slash burns on dry sites with shallow soil [45].  Fire can thus be used to control salal on dry sites but is often ineffective on wet sites [57].  Slash burning in Douglas-fir plantations of eastern Vancouver Island reduced the height and cover of salal while improving the nutrient status of Douglas-fir [131].  Often the moderate fires that reduce salal produce a positive response in conifer seedlings.  Where slash burning is contemplated, plots should be burned immediately after timber harvest for best results.  Because slash burns delay but do not eliminate salal, it is important that sites are planted within 2 years after logging and fire [16]. While slash burns often aid conifer regeneration, in some locations salal cover is not significantly reduced and competition remains a considerable problem.  Factors such as site characteristics, community composition, and fire intensity and severity are all important influences.  In old growth Douglas-fir forests of the western Cascades, salal may triple in cover during the first 5 years after logging and slash burn as shown below [31]:           1962       1963     1964     1965     1966     1967      1968          before     1st yr.   1st yr.            logging    after     after                     logging   slash burn %cover     5.9       1.1       0.5     1.3      1.6       2.2       3.0 %freq.    20.2       5.8       4.0     5.8      6.4       7.7       9.5 Response of salal after timber harvest and subsequent slash burns has been examined by a number of researchers [30,31,70,71,97,119,122,131, 140].

FIRE CASE STUDIES

SPECIES: Gaultheria shallon

1st CASE STUDY:
FIRE CASE STUDY CITATION : Tirmenstein, D., compiler. 1990. Salal response to clearcutting and spring or fall burning in a Douglas-fir-western redcedar-western hemlock forest, western British Columbia. In: Gaultheria shallon. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ []. REFERENCE : Lafferty, R. R. 1972. Regeneration and plant succession as related to fire intensity on clear-cut logged areas in coastal cedar-hemlock type: an interim report.  Internal Report BC-33. Victoria, BC: Department of the Environment, Canadian Forestry Service, Pacific Forest Research Centre. 129 p. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Lab, Missoula, MT. [82]. SEASON/SEVERITY CLASSIFICATION : Plot 6 - May 22, 1969/high Plot 7 - September 9, 1968/moderate STUDY LOCATION : The study site was located approximately 33 miles (53 km) east of Vancouver and 14 miles (22 km) north of Mission City, British Columbia. PREFIRE VEGETATIVE COMMUNITY : Douglas-fir (Pseudotsuga menziesii) dominated the overstory with scattered western redcedar (Thuja plicata) and western white pine (Pinus monticola) on the south and west aspects and western hemlock (Tsuga heterophylla) and western redcedar on the north and east aspects.  Red elderberry (Sambucus racemosa), willow (Salix spp.), mountain ash (Sorbus sitchensis), vine maple (Acer circinatum), red and ovalleaf huckleberry (Vaccinium parvifolium, V. ovalifolium), trailing blackberry (Rubus ursinus), thimbleberry (R. parviflorus), salal, fireweed (Epilobium angustifolium), dwarf Oregon grape (Mahonia nervosa), northern twinflower (Linnaea borealis), deerfern (Blechnum spicant), and various mosses were common in the preburn community. TARGET SPECIES PHENOLOGICAL STATE : not reported. SITE DESCRIPTION : Elevation:  500 feet (152 m). Parent materials:  bedrock was composed of quartz diorite and diorite,                    overlain with glacial till, outwash and minor                    lacustrine and aeolian deposits. Soils:  mixture of colluvium, loess, and ablation till, loamy with mixed         gravel throughout.  Climate:  marine and cool, no distinct dry         season.  An average of 203 frost-free days per year. FIRE DESCRIPTION :       rate of           residence         total fuel        energy released       spread            time              loading           (cal./m sq. x       (ft./min.)        (min.)            (g/m sq.)         1,000) Plot 6    22               85             15,840            22,709 Plot 7    15               50             30,308            45,799       initial duff            residual duff           % duff reduction       wt. (g/m sq.)           wt. (g/m sq.)           by weight Plot 6     6,700                  3,750                     44 Plot 7    10,000                  6,710                     33                      avg. initial      avg. initial      fuel consumed     total energy       fuel  (g/m sq.    loading (g/m      (g/m sq. x        (cal/m sq. x       x 1,000)          sq. x 1,000)      1,000)            1,000,000) Plot 6      8.322          15.022           3.058           22.709 x 10 Plot 7     20.321          30.308           7.946           45.799 x 10                   percent moisture - slash fuel component             fine fuel         med. fuel         large fuel             (.04-2.5 in.)     (.43-3.9 in.)      (4.0 in. or >)             (.01-1.0 cm)      (.1.1-10 cm)       (10.1 cm or >)    Plot 6      12.5               6.6                40.3      Plot 7      17.4              16.3                21.4                  percent moisture - organic fuel component             litter            fermentation            humus Plot 6      21.8                 102.7                120.8 Plot 7      11.8                 146.1                197.8 FIRE EFFECTS ON TARGET SPECIES : Plot 6 - high intensity burn:  Very few plants were present after the burn.  By the third growing season, small colonies of salal were scattered throughout the plot. Plot 7 - moderate intensity burn:  Salal was observed the first year after fire.  Salal continued to increase in abundance during the first three growing seasons after fire.  Recovery was as follows:                    1969        1970        1971                  postburn % freq.            51.1       58.0          58.0 % cover             1.1        2.9          5.6 FIRE MANAGEMENT IMPLICATIONS : High intensity burns are more effective in delaying the recovery of salal than moderate intensity burns.
2nd CASE STUDY:
FIRE CASE STUDY CITATION : Tirmenstein, D., compiler. 1990. Effects of slash burning on salal on eastern Vancouver Island, British Columbia. In: Gaultheria shallon. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ []. REFERENCE : Vihnanek, R. E.; Ballard, T. M. 1988. Slashburning effects on stocking, growth, and nutrition of young Douglas-fir plantations in salal-dominated ecosystems of east Vancouver Island.  Canadian Journal of Forest Research. 18: 718-722. [131]. SEASON/SEVERITY CLASSIFICATION : not reported/low to high STUDY LOCATION : The study site was located on the east side of Vancouver Island, British Columbia. PREFIRE VEGETATIVE COMMUNITY : Sites are in the wet and dry, coastal western hemlock (Tsuga heterophylla) subzone.  The understory was dominated by salal.  Each site supported planted 5- to 10-year-old Douglas-fir (Pseudotsuga menziesii), some of which had been burned. TARGET SPECIES PHENOLOGICAL STATE : not reported. SITE DESCRIPTION : Soils:  Brunisols or podzols developed in till, overlying volcanic or         sedimentary bedrock.  Slope:  0 to 60 percent.  Elevation:  1,650 to 2,650 feet (500-800 m).  Climate:  average annual water deficit - 4.2 to 5.2 inches (106-133 mm).           mean annual temperature - 41 to 47 degrees F (5.4-8.7 degrees           C). FIRE DESCRIPTION : Fire severity was estimated on the basis of remaining fuels and percent exposed mineral soil.  Fire severity ranged from low to high and was defined as follows: high - absence of all fine and most medium (3-9.5 cm diameter) fuels,       considerable consumption of large fuels and stumps and a large       difference in percent mineral soil exposure between paired burned and       unburned plots (15-60 percent). moderate - intermediate fuel characteristics, small to moderate difference in       paired mineral soil exposure (0-5 percent) between burned and unburned       plots. low - fine fuels present (< 2.5 cm in diameter), minimal charring of large       fuels (> 10 cm diameter) and stumps; small difference (0-4 percent) in       percent mineral soil exposed on burned and unburned areas. FIRE EFFECTS ON TARGET SPECIES : site burn       salal cover             height              exposed mineral #    sever-         (%)                 (cm)                soil (%)      ity*      burned   unburned    burned   unburned      burned  unburned 1    H           16        55         15        28           26       0 2    H           16        54         20        29           15       0 3    M           25        44         18        35            5       1 4    H            4        70         16        49           60       0 5    H            9        44         18        32           31       2 6    H            5        41         18        22           24       0 7    H            9        47         14        28           36       5 8    H            7        55         14        34           25       0 9    L           16        25         28        21            6      10 10   H            6        15         23        24           39      24 11   M           15        40         22        32            4       0 12   M           26        32         25        23            2       4 13   M           41        64         26        36            0       0 14   M           34        52         23        36            6       0 15   L           47        60         31        34            4       0 16   L           15        26         21        21            3       0 17   L           30        63         17        25            6       3 18   L           24        54         16        23            1       1 19   M           19        38         20        25           11       1 20   L           40        51         26        30            1       0  *L - low severity   M - moderate severity   H - high severity FIRE MANAGEMENT IMPLICATIONS : Salal cover and height growth can be significantly reduced by burning with corresponding increases in the height growth of Douglas-fir seedlings.  Vihnanek and Ballard [131] note that "results [of this study] suggest that slashburning should remain as a site preparation option in the dry salal-dominated forest ecosystems of eastern Vancouver Island.  However, it would be inappropriate to extrapolate the results of this study to other kinds of ecosystems."

References for species: Gaultheria shallon


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