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You are here >1Up Info > Wildlife, Animals, and Plants > Plant Species > Shrub > Species: Gaultheria shallon | Salal
 

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BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Gaultheria shallon | Salal
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]

Related categories for Species: Gaultheria shallon | Salal

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