Reclamation Suitability Criteria	Suitability Rating
Drought tolerance	Very high High Medium Low
Salt tolerance	X X
pH tolerance	X (acid)
Winter hardiness	X
Erosion control	X
Persistence	X
Palatability	X
Browse tolerance	X
Moisture preference	Moist to wet, well to poorly drained waterlogged soils
Soil preference	Wide texture range, includes peat soils

OTHER USES AND VALUES:

Willows are useful for erosion control and windbreaks [8], and provide medicine for many ailments including indigestion, worms, and stomach complaints [121]. It is the source of "diamond wood", used in carving and furniture making [256]. Scouler willow was used by Native Americans for traps, snares, sweathouse frames, burden baskets, toys, musical instruments, and medicine [203]. 

MANAGEMENT CONSIDERATIONS:

The deep root system, multiple stems, and ability to resprout from a subterranean root crown rather than surface resprouting may increase Scouler willow's tolerance to disturbance [24]. 

Fire can be used to maintain shrub productivity on seral brushfields and encourage shrub dominance after harvest. Tree establishment can be encouraged by excluding wildfire, using site preparation methods that do not encourage grass or shrub species, and planting and seeding pioneer species under the shade of killed shrubs [222].

Prescribed fire has been used to stimulate sprouting of Scouler willow that has grown out of reach of big game browsers [130,131,135]. In burned areas, the current annual growth of willows is considerably higher than in control areas, and fire increases both browse production and availability [18]. Prescribed fire was used in the 1960's to rejuvenate northern Idaho brushfields for elk winter range [136]. When prescribed burning causes greater than 50% canopy mortality, substantially higher current annual growth (CAG) of Scouler willow is produced. CAG of vigorously growing plants has a higher nutritive value and nutrient concentration than mature, slower growing plants [183]. Big game prefer browsing on the current annual growth of burned willows than that of unburned willows [18]. Utilization by elk demonstrated an obvious preference for the sprouts of burned Scouler willow following mechanical treatment and understory burning; the surviving plants were substantially more vigorous, with greater live biomass and better palatability than the unburned plants [15,16,23,131]. Scouler willow experiences a dramatic increase in available browse production immediately following burning, the result of shrub removal and resprouting that reduces the height and increases the availability of browse [130,134,139,140,263].  In one study, Scouler willow reached 10 feet in height 4 years after burning, with 80% of twig production available for elk browse. However, repeated treatment may be necessary to maintain browse availability [132]. Scouler willow browse production increased from 6.5 to 44.1 kg/ha 3 to 7 years after wildfire in Alaska.  The increase in production was due to an increase in the number of shrubs during the 1st 5 years and an increase in the number of twigs per shrub during all 7 years. Stems were available after the first 2 years; 4 years postfire, browse production in the burned area was twice that in the unburned area; and at 7 years postfire browse was 5 times greater. Browse production is likely to peak at 10-15 years and decrease by 20 years postfire [264].

Scouler willow responds well to both fall and spring burning; however, fall burning eliminates the following winter's food supply for animals. Spring burns regrow rapidly, with 5 foot (1.5 m) sprouts common by summer on burns in Idaho [130]. It has shown no decreased vigor from burning at 5 year intervals [135]. Brushfield burning to improve elk winter range in Idaho has often occurred during the spring or fall. However, another very important browse species, redstem ceanothus, is not always replaced by seedlings under these treatments because the seedcoat required high soil temperatures to crack and allow germination. Therefore, for winter range improvement, summer prescribed fires may need to be considered [259].

Willows produce, within limits, denser growth when they are browsed [8], with browsed stems branching laterally [253]. Browsing stimulates production of Scouler willow, though continuous browsing over several years may eventually deplete plant or soil reserves resulting in an eventual decline in productivity [263]. Following wildfires in Alaska, Scouler willow resprouts have suffered intense browsing by snowshoe hares, which often migrate to burned areas to feed. In some locations, 100% of the current annual growth of sprouts was removed for 2 succeeding winters [253,263]. This impact normally lasts for only a few years. Flowering and fruit production of Scouler willow are affected by heavy browsing, possibly resulting in low seedling success and recruitment, and browsing may suppress sprouting [24,34]. In burned and unburned clearcuts, grazed sites had no Scouler willow present, while ungrazed sites had 5 to 7% Scouler willow cover 11 years after treatment [61]. Where it is subject to overbrowsing, the loss of Scouler willow may result in substantial losses to elk herds [73].

Mechanical thinning may prevent suppression of Scouler willow in Douglas-fir/ponderosa pine forests [87]. Scouler willow increased in biomass and vigor in response to thinning in a Douglas-fir/ponderosa pine stand in Montana, indicating a positive response to overstory reduction [24]. Winter carrying capacity for mule deer may be substantially increased after mechanical treatment due to the establishment and growth of Scouler willow [87]. It is difficult to achieve control of Scouler willow using mechanical treatment due to the resprouting of damaged shrubs, and clearcutting can result in major canopy increases. Chemical treatments may be required if managing for ponderosa pine establishment due to its inability to compete with Scouler willow [227,228].

Herbicide treatments of Scouler willow aimed at increasing sprouting and maintaining production at a height available to big game have erratic and complex results;  the production of sprouts did not persist, but the reduction in crown height did persist [143]. Scouler willow is highly sensitive to light applications of 2,4-D and 2,4,5-T. With 2,4-D, basal sprouting was proportional to crown kill; the maximum crown height was reduced and the proportion of live crown within reach of deer and elk increased accordingly. Sprout production was most favored by treatment in August and least by treatment in October; plant kill was highest following September treatment [173]. In other studies, crown volume and cover of Scouler willow were reduced by 50% after herbicide treatment [157]. Herbicide control of Scouler willow may be effective in allowing Douglas-fir to regenerate in the seral brushfields of Idaho [92]. Percent control of Scouler willow with different herbicides ranges from 33 to 100% [158,159,160,161].

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BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Salix scouleriana | Scouler Willow

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• GENERAL BOTANICAL CHARACTERISTICS
• RAUNKIAER LIFE FORM
• REGENERATION PROCESSES
• SITE CHARACTERISTICS
• SUCCESSIONAL STATUS
• SEASONAL DEVELOPMENT

GENERAL BOTANICAL CHARACTERISTICS:

Scouler willow is generally a shrub, reaching 6 to 35 feet (2-10.6 m) in height [8,41,72,109,178,179,204].  It occasionally grows as a tree and may reach 65 feet (20 m) in height [8,32,34,53,76,171,178]. It is nonrhizomatous [218] with a deep, extensive root system [23,24,213,234] and may be multistemmed [23,24,79,204] or have one main trunk with twigs spreading or ascending [53]. It occurs in thickets and forests [202] forming a tall shrub layer in young stands [7,40], but is intolerant of shade and can persist only under thin canopies [7]. Beneath a tree canopy, Scouler willow exhibits a tall, upright growth form [72,218,234], but if top-killed by disturbance it resprouts from the root crown creating a round growth form up to 16 feet (4.8 m) in diameter [234].

Scouler willow has slender stems and branches [72] reaching 4 to 8 inches (10-20 cm) in diameter at the base [9,33,80]; smooth to flaky bark; and glabrous twigs [72]. Leaves are glabrous to leathery, obovate to oblanceolate, and occasionally serrate [9,54,72,109,121,194]. Twigs and leaf undersides of Scouler willow are densely to thinly pubescent, with appressed, reddish hairs [8,10,33,72,121]. Young twigs and vigorous shoots are often densely pubescent, while older stems are smooth [9,121,194]. Stripped bark of Scouler willow has a skunky odor [33,54,121].

A dioecious plant, Scouler willow has large, single-scaled, floral winter buds [10,32,89], and lacks a terminal bud [89]. Aments, expanding before or with leaves and quickly deciduous [32,121], are usually sessile or borne on a short spur shoot and flower profusely [9,10]. Fruiting catkins are 0.8 to 2.4 inches long (2-6 cm) by 0.4 to 0.6 inches thick (1-1.5 cm) with dark floral bracts 4-5 mm long, one gland, and a capsule 5-8 mm long with dense, short hairs and a somewhat long beak [32,33,54,121,194].

RAUNKIAER [197] LIFE FORM:

Phanerophyte

REGENERATION PROCESSES:

Scouler willow regenerates from both from seed and vegetatively. Scouler willow probably begins producing seed before 10 years of age [32]. Insects, especially bees, are important pollinators [88]. Seeds disperse in late spring, disseminated by wind and water [9,32,51,91,179,195,218]. These seeds have cottony hairs that allow them to travel long distances [32]. Seedlings may regenerate from windborne seed from as far as several miles away [87]. Scouler willow seeds are nondormant and remain viable for only a few days without moisture [227]. Willow seeds are characterized by a short seed life and rapid germination [179,187], and Scouler willow seeds usually germinate within 12 to 24 hours of dispersal [32]. The seeds are scarified by light burning [110] and moist mineral soil is required for germination and seedling establishment [187,218,227,267]. Zasada and others [267] found that on artificially seeded sites Scouler willow had substantially more germinants, 1st year survivors, and 3rd year survivors on heavily burned sites than moderately burned sites. 

In the laboratory, the germinative capacity of fresh seeds is high; normally 95 to 100% of seed germinates within 3 days [32,52]. The seeds contain substantial amounts of chlorophyll, and photosynthesis generally occurs as soon as the seed is moistened [32]. Field experiments found that Scouler willow seeds were not viable under dry outdoor conditions; most seeds did not germinate and those that did produced abnormal seedlings [52]. Light is required for good germination. Seed may be stored up to 4 to 6 weeks if kept moist at 32 to 41 degrees Fahrenheit (0-5 ^oC) [32]. 

High seed to seedling ratios on seeded plots suggest that seeding is an inefficient way of using seed, and planting artificially regenerated plants may be a more successful method of establishment and a more efficient use of available seed [267].

Scouler willow regenerates asexually by vigorously resprouting from a subterranean root crown [1,2,3,17,23,24,40,195]. This basal sprouting occurs in response to disturbance, including fire, flooding, and mechanical damage [1,2,47]. Scouler willow sprouts typically have a tall, fast growth response [169].

Scouler willow generally propagates readily from cuttings, with 40 to 80% rooting success [58,100,213] and roots developing within 4 weeks of planting in lab and field experiments [100]. Initial roots develop on the basal portion of the cutting [49] and continued rooting progresses along buried stem surfaces [62,100]. Root development is more rapid and successful with cuttings collected during the growing season [49,89]. Densmore and Zasada [49] found that cuttings planted in the field had a survival rate of 17% after two growing seasons. In general, willow cuttings are better able to establish if planted as rooted stock [168]. Softwood cuttings of Scouler willow root as well or better than hardwood cuttings, which may offer alternatives for vegetative propagation and flexibility in producing stock for conservation planting [62].

SITE CHARACTERISTICS:

Scouler willow has a wide range of adaptation [39]. It is found in drier habitats than most willows [72], occurring as scattered individuals on dry uplands as well as swamps, and mountain streams [8,33,107,108,121,129,179,200], and is capable of establishing in dry rocky conditions at high elevations [39]. Scouler willow commonly grows on gentle to moderate slopes [40,41,128]. While it does occur in riparian areas, Scouler willow is more common on upland sites above riparian areas [80,90,146,195], and is found primarily in forests, meadows, on slopes [54,193], and in transitional zones between riparian and upland areas [149]. 

Scouler willow may be found rarely on wet areas, but more commonly on moist areas or intermittent watercourses, and establishes both within gullies and at their bases [20,63,81,120,129]. In riparian areas, Scouler willow establishes on relatively stable banks and lower sideslopes in valleys, reflecting a preferred environment of transport rather than retention of materials and moisture [63,212].

Scouler willow is found on shallow to moderately deep soils [19,23,40,166]. It exhibits tolerance to a range of soil moisture conditions [38], occurring on moderately- to well-drained soils  [38,41,54,80,166,180,185,193].  Scouler willow commonly occurs on stony and silty soil with upper soil horizons dry during most of the growing season [63], and Forsyth [70] found that the cover of Scouler willow was greater at intermediate soil moisture levels than at either extreme. Soils on Scouler willow sites ranges from fine textured to gravelly [29,38,90,155,185,194,204]. It is commonly found on a variety of depositional land forms, including gravel bars [67]; glacial till; morainal blankets; river terraces; pumice flows; and alluvial, colluvial, and lacustrine deposits [7,55,152,155,185,189]. Parent materials are derived from a broad range of materials [19,23,174], but appear to be of little significance in distribution [125]. Scarification of the soil surface provides mineral soil important for Scouler willow establishment [87].

The elevational range of Scouler willow includes sea level to over 10,000 feet (3000 m) [4,8,10,11,19,20,23,29,33,42,51,53,54,60,63,72,129,149,174,205,244,249,260]. Scouler willow is found in the lower and upper montane elevational belts in the northwest U.S. [4,33,51,96,260] and higher elevations in the southwest [10,129,149,244,249]. Annual precipitation on these sites may range from 9.5 to 63 inches (240-1600 mm) [4,7,19,23,29,42].

In British Columbia, Scouler willow prefers drier, low elevation sites; mid- to lower slopes with rolling terrain and level to moderately sloping [90,194]. In the western U.S., it is most common in upland forests, cut-over areas, and burned areas in drier locations [53]. The range of Scouler willow stretches from valley bottoms to the lower subalpine forest zone in Idaho. At the low to mid-elevations it grows in moist riparian habitats and generally attains a small tree stature; at the higher elevations it becomes a medium to large shrub and tends to inhabit relatively drier sites [33]. It may occur more frequently on north- and east-facing slopes in the western U.S. [72,125,129,174]; Irwin and Peek [106] found that maximum height growth of Scouler willow in Idaho shrubfields occurred on east facing slopes. However, Mueggler [171] found that frequency and cover were not substantially related to aspect. Quaking aspen/Scouler willow communities in the Intermountain Region range from 5,800 to 7,400 feet (1,800-2,300 m) in elevation [174]. Distribution of Scouler willow in the southern part of its range may be somewhat restricted to montane riparian zones or other moist sites at high elevations [101,112,149,204,205,249], though in the southwest, Scouler willow occurs in mixed conifer forests on steep relatively dry slopes [10] and in low elevation canyons [113]. In California it may also grow from near sea level to 10,000 feet (3048 m) [204]. In northern Mexico, it has been found in the mountains on north-facing aspects [11].

Scouler willow is common in open areas following disturbance [165,178] and readily resprouts in ravines and on roadsides [47]. Its limited shade tolerance, tall growth habit, and sprouting ability enable it to persist in small openings on timbered sites [171,218,226]. Scouler willow occurrence increases with full sunlight [70,172]; Mueggler [171] found that the frequency and crown cover of Scouler willow was substantially higher under tree cover less than 25% than under tree cover exceeding 41%. Hungerford [103] found that maximum height of Scouler willow coincides with 40 to 50% of available light during canopy development. As a result of logging and slash burning in Douglas-fir/ponderosa pine forests in Montana, Scouler willow became established in direct proportion to the amount of stand opening and ground disturbance [53].

SUCCESSIONAL STATUS:

Scouler willow is a shade intolerant, persistent seral species [17,55,104,238]. It is often a minor understory component in a variety of forest types [5,43,44,45,59,68,72,76,77,85,95,110,163,185,258,260], occurring as scattered individuals in small openings. However, it increases after disturbance, including clearcutting, prescribed fire, soil disturbance, and wildfire [1,5,9,17,47,53,61,70,71,72,89,106,138,145,150,171,194,222]. 

Scouler willow is an early to mid-seral species [43,44,73,84,85,94,102,103,127,132,145,184,185,198,218,236]. Where not already present, it rapidly invades disturbed sites [24,45,91,214,238] facilitated by its wind-dispersed seed [216,265], or resprouts following canopy removal [1,69,265]. It capitalizes on moderate to severely burned sites [110,177]. In clearcuts and young stands it forms a tall shrub layer, comprising a substantial percentage of the plant cover [7,41,104]. In some areas, Scouler willow may dominate early seral plant communities following fire or clearcutting [28,55,150,210,211,265] and on river terraces and gravel bars [1,67]. It forms a mid-seral shrub layer, the Scouler willow layer group, in several habitat types of the northern Rocky Mountains [17,226,227,228,229,231]. In the past, these layer groups formed in response to stand-replacing wildfires, but may also develop following mechanical scarification in clearcuts. Scouler willow may persist in late seral and climax stands, but the layer groups fade as succession progresses [226,229].

Scouler willow reaches its highest frequency and cover in stands at least 20 years old, with maximum frequency and cover reached between 30 and 40 years [171]. Slight shade tolerance, tall growth habit, and ability to resprout allow Scouler willow to persist under moderately dense tree cover because small openings in the canopy can stimulate sprouting and rejuvenate individuals, making it somewhat less vulnerable to successional replacement [218,222,227]. It may remain present but substantially less abundant in climax cover types [17,41,55,68,76,95,185,260].  On paired stands of uncut and clearcut grand fir forest, Scouler willow appeared only in the clearcut stands, 7 to 16 years after disturbance [6,7]. Following a stand replacing fire, Scouler willow appeared in stands 30 to 90 years old but was not present in stands greater than 150 years old [7]. Presence and cover of Scouler willow has been found to decrease with increasing stand maturity [17,172,219,240]; in one study, both values decreased significantly (p<0.05) as stands progressed from immature (<90 years old) to old growth (>150 years old) [5]. The disappearance of Scouler willow from maturing stands is attributed to the increasing competition for light and moisture as the tree cover develops [34].

Successional trends in northern Idaho, following the removal of climax coniferous forests from wildfire or logging (occasionally accompanied by prescribed fire), show that the initial postfire vegetation is dominated by a grass-forb stage [42,72,92,115,133]. Within a few years this gives way to dense brushfields, of which Scouler willow is often a dominant or co-dominant [42,56,70,72,92,97,106,108,115,133,240]. These brushfields eventually return to conifer-dominated sites; the time frame depends upon fire intensity, reburn history, seed sources, climatic conditions, and site characteristics [72].

SEASONAL DEVELOPMENT:

Scouler willow bud development begins in April, leafing out occurs in April and May, stem elongation occurs May through July, and leaf fall occurs in July through November [56,181]. Earlier leaf fall is correlated with limited moisture availability [56]. Flowering occurs in April through June, with fruit ripening and seed dispersal occurring in May through July [32,254].

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FIRE ECOLOGY

SPECIES: Salix scouleriana | Scouler Willow

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• FIRE ECOLOGY OR ADAPTATIONS
• POSTFIRE REGENERATION STRATEGY

FIRE ECOLOGY OR ADAPTATIONS:

Willows are greatly favored by fire in most habitats [89,148,253].  As a survivor and off-site colonizer [117,237,241,242,243], Scouler willow is abundant following fire [146] and has a moderate regeneration period [121]. It is adapted to fire by rapidly resprouting from the root crown [145,167,179,216], and establishes from seed on severely burned sites [179]. Wind dispersed seeds facilitate rapid recolonization of burned areas [216,217]. In a north-west Montana study Scouler willow was found on 80% of burned sites with no previous Scouler willow presence [237]. Stand replacing fires favor regeneration of Scouler willow [167], and good response from Scouler willow seedlings can be expected on sites where fire damage is thorough enough to expose mineral soil [87].  However, it is rarely present on sites where more than 50% of the prefire overstory remains [70]. 

Scouler willow layer groups are distinct shrub layers that occur in various habitat types and are created by stand replacing fires [218,227,228,229,231]. Severe wildfires expose patches of bare mineral soil, encouraging the development of Scouler willow shrub layers [229,231]. These layer groups may also develop in response to mechanical scarification in clearcuts and broadcast burns, especially where exposed soil was mounded to trap water behind the mounds, creating well-watered seedbeds of mineral soil [229,231,233].

Scouler willow is frequently a dominant or codominant in the persistent seral brushfields of northern Idaho. These brushfields are likely the result of dry weather patterns after canopy removal and repeated severe fires, which remove most large woody material, litter, and herbaceous fuels. Standing shrubs comprise most of the biomass, and these brushfields can burn in almost any season. If surface fuels are continuous and dry, spring fires spread readily. In the summer, brushfields are often hot and dry, and conditions are exacerbated where nighttime inversions occur. Hot, dry winds during drought conditions can drive severe fires through the shrub layer, with Scouler willow readily regenerating from seed and resprouting [222].

In interior Alaska, Foote [69] identified six community developmental stages following fire in black spruce forests. These are: 1) newly burned, 2) moss-herb, 3) tall shrub-sapling, 4) dense tree, 5) hardwood or mixed hardwood-spruce, and 6) black spruce. Arising from resprouts, Scouler willow can average up to a few thousand stems per hectare 1 year following wildfire depending on preburn density, and thus is an important part of the newly burned stage. It is then often dominant or co-dominant through the tall shrub-sapling stage of succession for about 30 years. It thereafter becomes less frequent, as larger trees outgrow and overtop it.

Resprouting occurs in response to overstory thinning [217] where Scouler willow is well-distributed and in need of rejuvenation [87]. Generally, fast spreading fires produce more willow sprouts than slow fires that can damage root crowns [222]. Density and canopy coverage frequently increase after fire because root crowns produce multiple sprouts [179]. Four years postfire in Alaska, Scouler willow presence was 4 times greater on burned sites than on adjacent unburned sites [253]. In northern Idaho, Scouler willow cover was much higher on burned clearcuts than on unburned clearcuts, particularly where there had been repeated fires over a 30 year period [172]. Postfire immature stands (<90 years) in Montana have significantly more (p<0.05) presence and percent cover of Scouler willow than old growth stands (>150 years) [5].  The increased presence of Scouler willow in Douglas-fir/ponderosa pine stands following elimination of frequent fires is likely the result of stand opening associated with logging [19]. Resprouting Scouler willow creates a round growth form up to 16 feet in diameter and may as a result promote reestablishment of shade tolerant species like Douglas-fir, in turn posing a greater risk of stand replacing fires and favoring growth of Scouler willow [234].

Without fire, closing conifer canopies lead to the deterioration of Scouler willow [84]. In dense second growth stands of sequoia in California, Scouler willow debris creates a fuel hazard; formerly abundant stands of Scouler willow grew in dense clones that were shaded out and killed, forming dense tangles of fuel for wildfire [28].

Fire regimes for plant communities and ecosystems in which Scouler willow occurs are summarized below. For further information regarding fire regimes and fire ecology of communities where Scouler willow is found, see the "Fire Ecology and Adaptations" section of the FEIS species summary for the plant community or ecosystem dominants listed below.

Community or Ecosystem	Dominant Species	Fire Return Interval Range (years)
silver fir-Douglas-fir	Abies amabilis-Pseudotsuga menziesii var. menziesii	> 200
grand fir	A. grandis	35-200 [14]
California montane chaparral	Ceanothus and/or Arctostaphylos spp.	50-100
Rocky Mountain juniper	Juniperus scopulorum	< 35 [188]
western larch	Larix occidentalis	25-100
Engelmann spruce-subalpine fir	Picea engelmannii-Abies lasiocarpa	35 to > 200 [14]
black spruce	P. mariana	35-200
conifer bog*	P. m.-Larix laricina	35-200 [57]
blue spruce*	P. pungens	35-200 [14]
pinyon-juniper	Pinus-Juniperus spp.	< 35 [188]
whitebark pine*	P. albicaulis	50-200 [14]
Rocky Mountain lodgepole pine*	P. contorta var. latifolia	25-300+ [13,14,201]
Sierra lodgepole pine*	P. c. var. murrayana	35-200
Jeffrey pine	P. jeffreyi	5-30
western white pine*	P. monticola	50-200
Pacific ponderosa pine*	P. ponderosa var. ponderosa	1-47
Rocky Mountain ponderosa pine*	P. p. var. scopulorum	2-10
Arizona pine	P. p. var. arizonica	2-10 [14]
aspen-birch	Populus tremuloides-Betula papyrifera	35-200 [57,255]
quaking aspen (west of the Great Plains)	P. t.	7-120 [14,83,156]
mountain grasslands	Pseudoroegneria spicata	3-40 (10**) [13,14]
Rocky Mountain Douglas-fir*	Pseudotsuga menziesii var. glauca	25-100 [14]
coastal Douglas-fir*	P. m. var. menziesii	40-240 [14,170,199]
California mixed evergreen	P. m. var. m.-Lithocarpus densiflorus-Arbutus menziesii	< 35 [14]
redwood	Sequoia sempervirens	5-200 [14,65,246]
western redcedar-western hemlock	Thuja plicata-Tsuga heterophylla	> 200
western hemlock-Sitka spruce	T. h.-Picea sitchensis	> 200
mountain hemlock*	T. mertensiana	35 to > 200 [14]

*fire return interval varies widely; trends in variation are noted in the species summary
**mean

POSTFIRE REGENERATION STRATEGY [241]:

Tall shrub, adventitious bud/root crown
Initial off-site colonizer (off-site, initial community)

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FIRE EFFECTS

SPECIES: Salix scouleriana | Scouler Willow

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• IMMEDIATE FIRE EFFECT ON PLANT
• DISCUSSION AND QUALIFICATION OF FIRE EFFECT
• PLANT RESPONSE TO FIRE
• DISCUSSION AND QUALIFICATION OF PLANT RESPONSE
• FIRE MANAGEMENT CONSIDERATIONS

IMMEDIATE FIRE EFFECT ON PLANT:

Scouler willow is sensitive to fire [121] and may be top-killed [234]. Scouler willow crown mortality following fire can vary from 0 to 100% depending on fire severity [183]. Severe fires that destroy the organic layer may result in 100% aboveground mortality [145]. However, even when aboveground plant parts are destroyed by fire, underground plant parts usually survive.

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:

Scouler willow may be resistant to fire, and there is a greater than 65% chance that 50% of the population on a site will survive or immediately reestablish after passage of a fire with an average flame length of 12 inches (30.5 cm) [195]. Mortality of Scouler willow to burning was low in different prescribed fire treatments in Montana ponderosa pine stands [24].

PLANT RESPONSE TO FIRE:

Sprouting: Willows sprout quickly after fire if depth of the burn in the soil is low to moderate [93]. When fire is intense enough to kill live foliage but does not kill the vascular cambium, Scouler willow experiences vigorous epicormic sprouting from the root crown [2,3,19,24,30,31,47,76,103,131,134,139,140,142,153,171,172,191,222,226,230,239,247,261]. Root crowns of Scouler willow are often so large that some buds always survive, except when disturbance is really severe [169]. New shoots have been observed growing within days of a fire [69], and rapid resprouting after burning results in low overall plant mortality [19]. Multiple sprouts result in increased Scouler willow density following fire [116]. Scouler willow has a high postfire response rating; the species population will regain its preburn frequency and cover in 5 years or less [195]. 

Scouler willow increases dramatically following a variety of burn intensities, especially on relatively moist sites [195]. Basal area after 3 years may exceed that on unburned sites [253]. Dramatic increases in volume occur over the first 15 years postfire [239]. Resprout height growth may be dramatic, reaching up to 10 feet (3 m) after the first growing season [46,131,259]. Within 3 years after burning, plant crowns can average over 11 feet (3.4 m) in height [131]. Following a prescribed summer burn in Idaho, nearly 80% of height growth of Scouler willow over a 7-year period was attained within 2 growing seasons [140]. Following a summer wildfire in northern Idaho, Scouler willow reached peak cover values within 8 years [240]. 

Scouler willow plants that experience severe canopy mortality apparently concentrate their nutrients into vigorous new growth more than plants which experience only light canopy mortality. Analysis of aboveground plant part mortality classes from controlled burns in Idaho revealed that Scouler willow plants which experience top-kill exhibit the most vigorous regrowth. Twig growth (length and weight) of Scouler willow following fire was 3 to 4 times greater on plants with greater than 50% canopy mortality than on plants with less than 50% canopy mortality [183].

Seeding: Scouler willow also has the potential to regenerate from off-site seed sources [30,31,46,86,91,103,172,222,226,239,247,258], and can establish in moist mineral soil postfire [30,31,66,76,222,226]. Sowing Scouler willow seeds on different severity burns in upland black spruce sites in Alaska showed that germination occurred only on moderately (organic layers partially consumed) and severely (ash layer present, organic material in soil consumed or nearly so to mineral soil) burned seedbeds. Severely burned sites had the best germination percentages and represented the only burn severity class where Scouler willow seedlings survived past 3 years [267]. Scouler willow establishes quickly, but the rate of cover development or increase is slow [237,238,240].

Germinating seed originating from off-site plants often raises Scouler willow frequency far above what would be expected from on-site surviving plants [140,145,240]. Stickney [240] observed that after a stand-replacing wildfire in northern Idaho, Scouler willow seedlings made up the majority of the shrub component of the vegetation. The importance of seedlings in the postfire community was similarly observed by Lyon [140]. He recorded the postfire density of Scouler willow plants for 7 years, summarized below (density = # of plants >18 inches (46 cm) tall per 1,000 ft²):

		Postfire year
	Prefire - 1963	1 - 1964	2 - 1965	3 - 1966	4- 1967	5 - 1968	6 - 1969	7 - 1970
Density	0.3	0.1	0.2	0.2	0.6	2.4	3.6	4.4

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:

No entry

FIRE MANAGEMENT CONSIDERATIONS:

Prescribed fire is widely used as a wildlife management tool to rejuvenate decadent willow stands and stimulate sprouting. In areas where Scouler willow is scattered through the vegetation and in low vigor due to overbrowsing, prescribed fires that kill aboveground plant parts and expose mineral soils are favorable. This allows existing shrubs to sprout and creates favorable seedbeds for establishment of Scouler willow. In Douglas-fir/pinegrass habitat types of Montana, burning during late summer or fall exposes 30 to 50% of mineral soil [87]. A quick, hot fire maximizes sprouting while slower, longer burns cause more extensive damage and reduce browse [89]. The deep root system and multistemmed growth of Scouler willow allows for higher tolerance to disturbance [19], and it establishes rapidly in clearcut and heavily burned areas [61,80]. Scouler willow is favored by conditions on burned areas; it is scarce on areas not subjected to fire but very abundant on broadcast burns [171]. However, broadcast burns do not always burn hot enough to duplicate the effects of severe wildfire and create an adequate seedbed for Scouler willow, which is favored by light soil scarification [150,218,226,227,228,229]. Competition may limit Scouler willow establishment; the frequency and percent cover of Scouler willow were significantly less (p<0.05 and p<0.01 respectively) on a burned, artificially seeded site than on a burned, unseeded site [208].

On sites in northern Idaho, Scouler willow had substantially higher cover on a 30-year-old burn than after any intensive silvicultural treatment (ranging from thinning to clearcut), with no presence in the control [104]. However, after 30 to 40 years of fire exclusion in ponderosa pine forests, Scouler willow presence increased [22]. Logging and fire suppression allowed vigorous development of Scouler willow in a Douglas-fir/ponderosa pine community [87]. 

Prescribed fire enhances vigorous regrowth and germination of Scouler willow, and it is effective in increasing biomass [24]. In the cedar-hemlock zone of Idaho, it generally produces the most cover of any shrub species on broadcast burned areas. Cover and frequency of Scouler willow is substantially higher on broadcast burns than on areas not broadcast burned, as are the mean height values [171]. Total shrub volume of Scouler willow in Douglas-fir stands increased from 15 to 80% 2 years postfire [139]. Mean canopy coverage of Scouler willow  increased significantly (p<0.01) following selective logging and low intensity broadcast burning (intensity ~ 127 kcal/m/s) [12]. In the 1st year following burning, Scouler willow may produce up to 28 times as many sprouts as the previous year [136,162]. Scouler willow may grow significantly wider and taller (p<0.05) after fall burning than after spring burning [136]. However, fall burning removes the following winter's food supply for animals, while spring burns have substantial regrowth by summer [130]. After stand mechanical treatment and understory burning, Scouler willow was reduced by 9% from mechanical damage and an additional 16% from fire. The surviving plants were substantially more vigorous post-treatment [15,16]. While modest Scouler willow mortality may result after overstory removal and prescribed fire, the percentage of high vigor plants increases; in one study the amount of vigorous plants increased from 15% at pretreatment to 70% post treatment [23]. In northern Idaho, burning at 5-year intervals did not  result in decreased vigor [135]. Scouler willow was not markedly affected by burning until it suffered deep charring of the root crown. The following table presents the change in Scouler willow population characteristics in response to different treatments (% change is relative to pretreatment conditions) [23]:

Treatment	Cover reduction	Mortality	High vigor plants
Control	1	3	15
Harvest only (shelterwood cut)	33	14	60
Low consumption (shelterwood cut and low consumption burn)	62	22	71
High consumption (shelterwood cut and high consumption burn)	58	26	69

Shelterwood cuts combined with prescribed burning in a ponderosa pine resulted in modest Scouler willow mortality; plants remaining in the harvest only and burned treatments had higher vigor than those in the control [19,23].

A summary of Scouler willow's response to different types of disturbance is presented below [228]:

Type of disturbance:	Clearcut, no site prep	Shelterwood cut, mechanical scarification	Clearcut, mechanical scarification	Clearcut, broadcast burn	Stand destroying wildfire
Response:	major vegetative response	minor vegetative response	major vegetative response, minor response from seed	major vegetative response, minor response from seed	major vegetative response, minor response from seed

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Salix scouleriana: References

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Salix scouleriana Index

Related categories for SPECIES: Salix scouleriana | Scouler Willow

• Introductory
• Distribution and Occurrence
• Value and Use
• Botanical and Ecological Characteristics
• Fire Ecology
• Fire Effects
• References
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