Index of Species Information

SPECIES:  Picea engelmannii

Introductory

SPECIES: Picea engelmannii
AUTHORSHIP AND CITATION : Uchytil, Ronald J. 1991. Picea engelmannii. 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 : PICENG SYNONYMS : Abies engelmannii Parry Picea engelmannii var. glabra Goodman Picea glauca var. engelmannii (Parry) Boivin Picea glauca ssp. engelmannii (Parry) T. M. C. Taylor Picea columbiana Lemmon SCS PLANT CODE : PIEN COMMON NAMES : Engelmann spruce Columbian spruce mountain spruce TAXONOMY : The genus Picea consists of about 30 species of evergreen trees found in cool, temperate regions of the northern hemisphere. Seven species of Picea, including Engelmann spruce, are native to North America. The currently accepted scientific name of Engelmann spruce is Picea engelmannii Parry ex Engelm. [45,53]. There are no recognized races or geographic varieties. Natural hybridization between species of Picea is common. Engelmann spruce x white spruce (Picea glauca) hybrids are common where the ranges of these species overlap. Natural crosses between these species occur from central British Columbia as far south as eastern Washington and Yellowstone National Park [23]. Within this area, trees at low elevations closely resemble pure white spruce. Pure Engelmann spruce tends to dominate at higher elevations [23]. Engelmann spruce x white spruce hybrids are common throughout low elevations in British Columbia [32]. Using artificial pollination techniques, Engelmann spruce has been successfully crossed with white spruce, blue spruce (P. pungens), and Sitka spruce (P. sitchensis) [32]. LIFE FORM : Tree FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY

DISTRIBUTION AND OCCURRENCE

SPECIES: Picea engelmannii
GENERAL DISTRIBUTION : Engelmann spruce is widely distributed throughout the mountains of the western United States and Canada. It occurs from central British Columbia and Alberta as far south as New Mexico and Arizona [6]. It is cultivated in Hawaii [101]. In the Pacific Coast region, Engelmann spruce is only a minor component of high-elevation forests. It grows from the Coastal Range in west-central British Columbia, south along the east slope of the Cascades through Washington and Oregon to Mount Shasta in northern California [6]. In the Rocky Mountains Engelmann spruce is a major component of high-elevation forests. It grows from southwestern Alberta, south through the mountains of eastern Washington, Idaho, and western Montana to the high mountains of southern Arizona and New Mexico [6]. ECOSYSTEMS : FRES20 Douglas-fir FRES22 Western white pine FRES23 Fir - spruce FRES24 Hemlock - Sitka spruce FRES25 Larch FRES26 Lodgepole pine FRES28 Western hardwoods FRES44 Alpine STATES : AZ CA CO HI ID MT NV NM OR UT WA WY AB BC BLM PHYSIOGRAPHIC REGIONS : 2 Cascade Mountains 4 Sierra Mountains 5 Columbia Plateau 6 Upper Basin and Range 7 Lower Basin and Range 8 Northern Rocky Mountains 9 Middle Rocky Mountains 11 Southern Rocky Mountains 12 Colorado Plateau 16 Upper Missouri Basin and Broken Lands KUCHLER PLANT ASSOCIATIONS : K004 Fir - hemlock forest K012 Douglas-fir forest K014 Grand fir - Douglas-fir forest K015 Western spruce - fir forest K018 Pine - Douglas-fir forest K020 Spruce - fir - Douglas-fir forest K021 Southwestern spruce - fir forest K052 Alpine meadows and barren SAF COVER TYPES : 201 White spruce 203 Balsam poplar 205 Mountain hemlock 206 Engelmann spruce - subalpine fir 208 Whitebark pine 209 Bristlecone pine 210 Interior Douglas-fir 212 Western larch 213 Grand fir 215 Western white pine 216 Blue spruce 217 Aspen 218 Lodgepole pine 219 Limber pine 224 Western hemlock 226 Coastal true fir - hemlock 227 Western redcedar - western hemlock 228 Western redcedar SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Climax stands consisting entirely of Engelmann spruce are somewhat scattered and often restricted to wet or cold habitats [4,88]. The Engelmann spruce series is generally recognized by the absence or scant representation of subalpine fir. This is because many ecologists, especially those working in the northern Rocky Mountains, believe that only in the absence of subalpine fir does Engelmann spruce dominate at climax [22,67,98]. Throughout the Rocky Mountains, Engelmann spruce occurs in widespread forests within the subalpine fir or white fir (Abies concolor) habitat-type series either as a climax codominant or long-lived seral species [4,98]. In the northern Rocky Mountains, Engelmann spruce is considered a long-lived seral species in fir habitat types. In the central and southern Rocky Mountains, Engelmann spruce and subalpine fir often codominate at climax; however, these forests are classified under the subalpine fir series to be consistent with habitat-type usage elsewhere [40,47]. Published classification schemes listing Engelmann spruce as an indicator or dominant in habitat types (hts), community types (cts), plant associations (pas), ecosystem associations (eas), site types (sts), riparian zone associations (rzas), or dominance types (dts) are presented below: Area Classification Authority AZ, NM: ----- forest & woodland hts Layser & Schubert 1979 ----- forest hts Moir & Ludwig 1979 Apache, Gila, Cibola NFs forest hts Fitzhugh & others 1987 s of Mogollon n AZ: San Fransisco general veg. cts, hts Rominger & Paulik 1983 n AZ, n NM forest hts Larson & Moir 1987 CO: Arapaho & Roosevelt NFs forest hts Hess & Alexander 1986 Gunnison & Uncompahgre NFs forest hts Komarkova & others 1988 Routt NF forest hts Hoffman & Alexander 1980 White River NF forest hts Hoffman & Alexander 1983 w CO riparian pas Baker 1989a ID: Sawtooth, White Cloud, Boulder, & Pioneer Mtns general veg. cts Schlatterer 1972 c ID forest hts Steele & others 1981 n ID forest hts Cooper & others 1987 e ID, w WY forest hts Steele & others 1983 riparian cts Youngblood & others 1985a MT forest hts Pfister & others 1977 riparian dts Hansen & others 1988 c, e MT riparian cts, hts Hansen & others 1990 nw MT riparian hts, cts Boggs & others 1990 sw MT riparian sts, cts, hts Hansen & others 1989 NM: Cibola NF forest hts Alexander & others 1987 Lincoln NF forest hts Alexander & others 1984 n NM, s CO forest hts Develice & others 1986 OR: Wallowa-Whitman NF steppe & forest pas Johnson & Simon 1987 Deschutes, Ochoco, Fremont & Winema NF's riparian rzas Kovalchik 1987 OR, WA: Blue Mtns forest & nonforest cts Hall 1973 c, s UT forest hts Youngblood & Mauk 1985 n UT forest hts Mauk & Henderson 1984 UT, se ID riparian cts Padgett & others 1989 WA: Okanogan NF forest pas Williams & Lillybridge 1983 WY: ----- riparian cts Olson & Gerhart 1982 Medicine Bow NF forest hts Alexander & others 1986 Bighorn Mtns forest hts Hoffman & Alexander 1976 Wind River Mtns forest hts Reed 1976 USFS R-4 aspen cts Mueggler 1988 AB general veg. cts Moss 1955 w-c AB forest cts Corns 1983 general veg. eas Corns & Annas 1986 BC: Prince Rupert Forest Region general veg. eas Pojar & others 1984

MANAGEMENT CONSIDERATIONS

SPECIES: Picea engelmannii
WOOD PRODUCTS VALUE : Engelmann spruce is an important commercial wood in the United States. The wood is white, odorless, lightweight, straight grained, soft, stiff, and can be readily air dried. It is easy to work, glues well, holds nails fairly well but has only average paint-holding properties. The wood is primarily used for lumber for home construction and for prefabricated wood products. Less common uses include veneer in plywood manufacture, poles, and specialty items, such as food containers, violins, pianos, and aircraft parts. Spruce has not been used much for pulp and paper, although its pulping properties are excellent [6]. IMPORTANCE TO LIVESTOCK AND WILDLIFE : Livestock: Livestock generally do not browse Engelmann spruce [4,89]. Wildlife habitat: Engelmann spruce-subalpine fir forests provide forage and habitat for a wide variety of small and large wildlife species [4,7,57,58]. However, these properties are characteristic of where spruce grows and the understory species associated with it rather than to the species itself. Animals that inhabit Engelmann spruce stands include moose, elk, mule deer, woodland caribou, porcupine, snowshoe hare, red squirrel, chipmunks, and voles. A partial list of birds that nest and feed in Engelmann spruce trees includes the mountain chickadee, Williamson's sapsucker, red-breasted nuthatch, brown creeper, and owls and woodpeckers [82]. Wildlife food: The young growth of Engelmann spruce is occasionally browsed by ungulates, but it is not an important food item and is probably only taken as a last resort [4]. Spruce grouse and blue grouse may feed extensively on buds and needles [56,80]. Squirrels sometimes clip and eat twigs and buds [77]. Engelmann spruce seeds are eaten by several species of small mammals and birds. Red squirrels, chickarees, and chipmunks eat seeds from cached cones [6,98]. Engelmann spruce seeds are also eaten off the ground or snow by chipmunks, mice, and voles [4]. Numerous species of birds, including chickadees, nuthatches, crossbills, and the pine siskin, remove and eat seeds from spruce cones [36,56]. Small birds may make considerable use of spruce seeds, but their foraging is scattered and sporadic throughout subalpine forests [36]. PALATABILITY : The palatability of Engelmann spruce to livestock and big game is low [26]. The seeds are palatable to small mammals and birds. NUTRITIONAL VALUE : Engelmann spruce is low in protein but fair in energy value [26]. A study in Montana found the following concentration of elements in Engelmann spruce needles and twigs [86]: 1-yr-old green needles twigs<.25 inch in diameter Calcium (mg/g [mean]) 6809 4028 Copper 7 9 Iron 57 237 Potassium 6914 7034 Magnesium 810 747 Manganese 669 323 Nitrogen 10911 4621 Sodium 100 128 Phosphorus 1841 1264 Zinc 69 72 Ash (percent [mean]) 5 3 COVER VALUE : Big game: Engelmann spruce provides excellent hiding and thermal cover for deer, elk, moose, bighorn sheep, and bear [40,50,98]. Dense stands of this species can provide cool summertime shade for big game. High-elevation stands provide bedding sites and protection from storms for bighorn sheep, mule deer, and elk [59]. Small mammals and birds: Small Engelmann spruce trees provide good year-round hiding cover for small animals. Blue grouse, which overwinter in conifers at high elevations, use spruce trees for protective cover and roosting sites [50,80]. Spruce trees in the Engelmann spruce/soft leaved sedge (Carex disperma) habitat type in central Idaho provide important nesting sites for the MacGillivray's warbler, American robin, and warbling vireo [89]. Engelmann spruce snags are used by numerous cavity-nesting birds. Snags greater than 11 inches (28 cm) d.b.h. are most often used [82]. VALUE FOR REHABILITATION OF DISTURBED SITES : Engelmann spruce can be planted on disturbed sites within forest vegetation types where it naturally occurs. It is primarily used for reforestation projects on cool, moist sites below upper timberline. It has been used to a limited extent for revegetation and long-term stabilization of high-elevation mine spoils [4,96]. In west-central Alberta, Engelmann spruce x white spruce hybrids were observed invading coal mine spoils at high elevations [76]. Planting nursery stock is more successful than direct seeding. Most commonly, 2- or 3-year-old bareroot or container-grown stock is planted following snowmelt [4,96]. Since seedlings are sensitive to direct sunlight, they should be planted in the protective shade of stumps, logs, or vegetation [4]. Artificial shade also is effective in protecting seedlings from wind and sun [15]. Two- to 4-foot tall (0.6-1.5 m), open-grown Engelmann spruce seedlings dug from the wild before breaking dormancy have shown good survival when transplanted [15]. Methods for collecting, processing, testing, storing, and planting seed, and for care and transplanting of bareroot and container-grown Engelmann spruce seedlings have been described in the literature [4,96]. OTHER USES AND VALUES : Engelmann spruce is sometimes used as an ornamental landscape plant. It has been used for screenings, windbreaks, and as a specimen tree [90]. Native Americans used Engelmann spruce for numerous purposes. The bark was often peeled into sheets and used for making canoes, baskets, and roofing. The fibrous roots were used to make rope, and the boughs and needles to make incense, body scents, and cleansing agents. Various teas and poultices were made from Engelmann spruce for medicinal purposes. Native Americans occasionally ate the inner bark [92]. OTHER MANAGEMENT CONSIDERATIONS : Timber harvest: Clearcutting and group selection silvicultural methods favor Engelmann spruce over true firs (Abies) and hemlocks (Tsuga) but increase the proportion of intolerant associates such as lodgepole pine (Pinus contorta) and Douglas-fir (Pseudotsuga menziesii) [6]. Shelterwood and individual tree selection tend to favor more tolerant associates. The seed tree method is generally not used because of the susceptibility of Engelmann spruce to windthrow. In the Rocky Mountains, clearcutting and shelterwood cutting have been the most commonly used harvesting methods in old-growth Engelmann spruce-subalpine fir stands because these stands tend to be even aged and overmature [3]. Successful natural regeneration of Engelmann spruce following logging is usually accomplished through mechanical scarification or broadcast burns which expose at least 40 percent of the mineral-soil seedbed [3]. Silvicultural systems and cutting methods for managing Engelmann spruce are described in detail in the literature [3,6]. Disease: The most common disease of Engelmann spruce is caused by wood-rotting fungi which results in root or butt decay. Spruce broom rust is also common in spruce-fir forests and causes bole deformation and spike tops, increases susceptibility to wind breakage, and provides entry points for decay fungi in spruce [7]. Dwarf mistletoe (Arceuthobium microcarpum) causes heavy mortality of spruce trees in Arizona and New Mexico [7]. Insects: The spruce beetle is the most serious insect pest of Engelmann spruce. Outbreaks are associated with extensive windthrow because downed trees provide a good food supply, causing a rapid expansion of beetle populations. Unabated logging slash has been responsible for past outbreaks [7].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Picea engelmannii
GENERAL BOTANICAL CHARACTERISTICS : Engelmann spruce is a long-lived, native, coniferous, evergreen tree. It is one of the largest of the high-elevation mountain conifers. Mature trees have a narrow, pyramid form and short, compact branches. Within natural stands, mature trees average 15 to 30 inches (38-76 cm) in diameter; the average dominant height varies from 45 to 130 feet (14-40 m), depending on site quality and density. Larger individuals are not uncommon and may exceed 40 inches (102 cm) in diameter and 160 feet (49 m) in height. Engelmann spruce is long-lived; dominant trees are often 350 to 450 years old, and 500- to 600-year-old trees are not uncommon [7]. The crowns of trees within a stand normally make up 50 to 70 percent of the total height of the tree [4]. Dead lower limbs tend to be persistent. The crowns of open-grown trees often extend down to the ground. In alpine areas just above treeline, Engelmann spruce often forms a krummholz. At treeline in northern Idaho, mature Engelmann spruce generally do not exceed 65 feet (20 m) in height, and progressively become more stunted as elevation increases, forming krummholz at the most severe, high-elevation sites [17]. The four-sided, acute-tipped needles are not particularly sharp, are deep bluish-green, and are 0.8 to 1.2 inches (2-3 cm) long [21,41]. The young twigs are finely pubescent, a characteristic which differentiates this spruce from white spruce, which has glabrous twigs. The bark is very thin, grayish-brown on young trees but at maturity becomes purplish brown to russet and is broken into loosely attached scales. Engelmann spruce is generally shallow rooted, but laterals may penetrate to a depth of 8 feet (2.4 m) in deep, porous, well-drained soils [4]. Engelmann spruce is monoecious. Female cones are light brown, 1.5 to 2.4 inches (4-6 cm) long, and occur in the upper part of the crown [21]. Male cones are usually found lower in the crown than female cones. Engelmann spruce seeds are about 0.12 inch (3 mm) long and have a single, well-developed wing about twice as long as the seed [21,77]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : Cone and seed production: Engelmann spruce can begin producing cones when 15 to 40 years old and 4 or 5 feet (1.2-1.5 m) tall, but under closed forest conditions seed production is generally not significant until trees are older and taller [6]. Within natural stands, most seed is produced by dominant trees greater than 15 inches (38 cm) d.b.h. Although yearly production is erratic, Engelmann spruce is considered a moderate to good seed producer. Good to bumper crops are produced every 2 to 5 years [4,6]. The seeds are light, averaging 135,000 per pound (297,500/kg) [77]. Seed predation: Cone and seed insects, especially the spruce seed worm, reduce seed yields. Seed production in Colorado was reduced an average of 28 percent by insects over a 4-year period [4]. Small mammals consume considerable amounts of freshly fallen seed off the forest floor, but the extent of loss is not known [6]. Dispersal: Engelmann spruce seed is generally shed by the end of October, but some may continue to fall throughout the winter. The winged seeds are wind dispersed. Seeds travel primarily with the prevailing winds, but upslope drafts can influence dispersal at low and middle elevations. Seed is generally dispersed within 300 feet (91 m) of a windward source; when bumper seed crops occur, about 5 to 10 percent of the seed may be dispersed as far as 600 feet (183 m) [6,63]. Seed dispersed during winter can travel great distances by skidding over glazed snow [58]. Viability and germination: The viability of Engelmann spruce seed is rated as good [7]. Germination averages 69 percent, which is much higher than that of associated species [7,77]. Under natural conditions, seeds overwinter under snow and germinate 2 to 3 weeks following snowmelt [6,83]. Occasionally germination may occur after summer rains or be delayed until the second year [4]. Seedling establishment and survival: Seedlings establish best on mineral soil. Decayed wood, duff, and litter are poor seedbeds because they dry out rapidly [6]. In general, seedlings that establish on organic matter deeper than 2 inches (5 cm) soon die because their shallow roots cannot penetrate to mineral soil before the surface organic layer dries out [4]. At middle to upper elevations, seedling survival may be greater on duff because the duff helps protect seedlings from high-intensity summer rain storms and from frost heaving [29]. In the central and southern Rocky Mountains, seedlings do not establish well in the open. Intense light on open slopes can inhibit photosynthesis, which eventually kills the seedling [63]. Direct sunlight also dries out seedbeds. Seedlings survive best under conditions of shade, cool temperatures, and adequate soil moisture [6]. Engelmann spruce requires a mineral soil seedbed; subalpine fir, however, is able to establish in duff because of its rapid root growth. Consequently, subalpine fir seedlings usually outnumber spruce seedlings in the understory of spruce-fir stands, even where Engelmann spruce dominates the overstory. Thus, even though it is short-lived, many ecologists consider subalpine fir better able to regenerate under climax conditions than Engelmann spruce. Growth: Engelmann spruce seedlings grow very slowly. One-year-old seedlings are seldom over 1 inch (2.5 cm) tall, and 5-year-old seedlings are usually 1 to 4 inches (2.5-10 cm) tall. Ten-year-old seedlings may be only 6 to 8 inches (15-20 cm) tall under heavy shade and 10 to 12 inches (25-30 cm) tall under partial shade. Under a dense overstory, seedlings may be severely suppressed; 3- to 5-foot-tall trees may be 100 years old. In light shade or full sun trees may grow to 4 or 5 feet (1.2-1.5 m) in height in about 20 years [6]. Vegetative reproduction: Near timberline, where the species assumes a dwarf or prostrate form, Engelmann spruce frequently reproduces by layering. Layering also occurs when only a few trees survive fire or other disturbances, but once numbers have increased enough to improve germination and establishment, layering decreases. In closed forest stands layering is negligible [7]. SITE CHARACTERISTICS : Engelmann spruce is found in some of the highest and coldest forest environments in the western United States, characterized by long, cold winters with heavy snowpack and short, cool summers [7]. It extends down to lower elevations along stream bottoms where cold air flows down the valley and collects in localized frost pockets [22]. It is generally found on moist and cool sites, but at timberline it may occur on somewhat dry sites. At middle elevations, pure stands are usually found on alluvial terraces, wet benches, bottomlands, slopes with seeps or cold north or east aspects [67,88]. It occurs on all aspects at timberline. Stand condition and associated conifers: Engelmann spruce forms pure stands but is more commonly associated with subalpine fir. These species frequently occur as codominants forming widespread subalpine forests. In the central and southern Rocky Mountains, Engelmann spruce commonly makes up 70 percent of overstory trees, and subalpine fir dominates the understory. Within Engelmann spruce-subalpine fir forests in this region, the spruce tends to be more important at higher elevations and on wetter sites, while subalpine fir is more abundant on drier lower elevation sites [4]. In Montana, pure Engelmann spruce stands are often found in cool ravines at lower elevations than subalpine fir [67]. Other associated conifers, which vary by latitude and elevation, are listed below [6]: Location Elevation Associates northern Rocky Mtns low and western white pine (Pinus monticola), and Cascade Mtns middle western redcedar (Thuja plicata), western hemlock (Tsuga heterophylla), Douglas-fir, grand fir (Abies grandis), lodgepole pine high Pacific silver fir (A. amabilis), mountain hemlock (Tsuga mertensiana), subalpine larch (Larix lyallii), whitebark pine (Pinus albicaulis) central and southern low and lodgepole pine, Douglas-fir, blue Rocky Mtns middle spruce, white fir, aspen (Populus tremuloides) high corkbark fir (Abies lasiocarpa var. arizonica), bristlecone pine (Pinus aristata), limber pine (P. flexilis) Understory associates: Understory vegetation is extremely variable, changing with elevation, exposure, and soil moisture. Habitat type and plant association guides describe characteristic understory plants for differing sites. Soil: Engelmann spruce grows best on moderately deep, well-drained, loamy sands and silts, and silt and clay loam soils developed from volcanic lava flows and sedimentary rock. It also grows well on alluvial soils where the underlying water table is readily accessible. It grows poorly on shallow, dry, coarse-textured sands; gravels developed primarily from granitic and schistic rock; coarse sandstones and conglomerates; rocky glacial till; heavy clay surface soils; and saturated soils [6]. Elevation: Elevational ranges for Engelmann spruce are described below [6]: Cascade Mountains -- generally between 4,000 and 6,000 feet (1,219 and 1,829 m); at 8,000 feet (2,438 m) on sheltered slopes and at 2,000 feet (610 m) in cold pockets along streams and valley bottoms Rocky Mountains: ID, MT, adjacent mtns eastern WA and OR -- between 2,000 and 9,000 feet (610 and 2,743 m); above 6,000 to 7,500 feet (1,829-2,286 m) a minor component of the stand; below 5,000 feet (1,676 m) confined to moist, lower slopes and cold valley bottoms UT, WY, and CO -- generally 9,000 to 11,000 feet (2,743-3,354 m); as low as 8,000 feet (2,438 m) along cold stream bottoms and sometimes as high as 11,500 (3,506 m) AZ and NM; plateaus of s UT -- between 8,000 and 12,000 feet (2,438 and 3,658 m); most common between 9,500 and 11,000 feet (2,896 and 3,354 m) SUCCESSIONAL STATUS : In the Rocky Mountains north and south of Montana and Idaho, Engelmann spruce and subalpine fir often codominate at climax to form extensive Engelmann spruce-subalpine fir forests [4]. These spruce-fir forests are usually classified as subalpine fir climax series habitat types. In the understory of these stands, subalpine fir seedlings usually outnumber Engelmann spruce seedlings because they are more shade tolerant and readily establish on duff seedbeds. However, Engelmann spruce is longer lived and usually the largest tree in the stand. There is little evidence that Engelmann spruce will ever be replaced by subalpine fir in these regions [47]. In the Rocky Mountains of Montana and Idaho, and in the mountains of eastern Washington and eastern Oregon, Engelmann spruce is usually considered seral to subalpine fir. Subalpine fir may form pure stands at climax, but Engelmann spruce is also often present because it outlives subalpine fir and persists to climax [4]. In eastern Washington and northern Idaho, Engelmann spruce is seral to grand fir, western redcedar, and western hemlock [22]. However, in Montana, eastern and central Idaho, and western Wyoming, Engelmann spruce may attain climax dominance on the wettest habitat types where it appears more successful than subalpine fir [88]. Farther east, progressing away from the Pacific maritime influence, the importance of Engelmann spruce increases and that of subalpine fir decreases [88], and in parts of central and southwestern Montana, Engelmann spruce may be dominant on well-drained benches and droughty soils [67]. On sites where Engelmann spruce attains climax dominance or codominance, succession following disturbance may vary depending on the severity and type of disturbance, elevation, and availability of seed. Near treeline, it may take 100 years or more for Engelmann spruce to establish seedlings following fire because an increase in herbaceous species prevents seeds from reaching mineral soil, and the harsh climate kills many seedlings that do establish [12,85]. Within subalpine stands, Engelmann spruce may establish immediately following disturbance if mature trees survive to provide seeds, and seral species such as lodgepole pine and aspen are scarce. Aspen and lodgepole pine are the most common seral species and often dominate subalpine forests following fire [6,47,85]. These species grow rapidly and quickly overtop any Engelmann spruce seedlings that may establish at the same time. Aspen stands can sometimes persist for decades or even centuries when conifer seed trees are eliminated [25,89]. When lodgepole pine establishes immediately following stand-destroying fires, it often forms dense even-aged stands that dominate for 100 to 300 years. Because it is shade tolerant, Engelmann spruce eventually establishes under the pine canopy, usually within 100 years, and attains dominance as the pine stand begins to break up [24,54]. On some of the lower elevation Engelmann spruce and subalpine fir habitat types, Engelmann spruce will not achieve climax dominance or codominance because of repeated fires which favor shade-intolerant seral conifers. Many of these habitat types are in midsuccessional stages; Douglas-fir, lodgepole pine, western larch (Larix occidentalis), or limber pine dominate the overstory [67,88] SEASONAL DEVELOPMENT : Pollen is generally shed from late May to early June at lower elevations and from late June to early July at higher elevations. Cones develop rapidly and are full size by August. Cones open and seeds are shed in late September and October, but some continue to fall throughout the winter. After seed dispersal, most cones fall during the winter, but some may persist for longer periods [4]. Phenological observations of Engelmann spruce in Montana, Idaho, and Wyoming made from 1928 to 1937 are presented below [79]: Buds Pollen shed Winter Cones Cones burst begins ends buds formed full size open (east of Continental Divide in Montana, and in Yellowstone NP) avg date June 16 June 17 July 3 Aug 3 Aug 17 Aug 30 earliest May 21 May 18 May 30 June 26 July 19 Aug 18 latest July 14 July 3 July 20 Sept 17 Sept 5 Sept 21 (west of Continental Divide in western Montana and northern Idaho) avg date May 27 June 1 June 7 Aug 23 Aug 6 Sept 8 earliest May 5 Apr 26 May 12 June 14 June 20 Aug 11 latest July 10 June 11 May 12? Oct 11 Sept 24 Oct 5

FIRE ECOLOGY

SPECIES: Picea engelmannii
FIRE ECOLOGY OR ADAPTATIONS : Plant adaptations to fire: Engelmann spruce is very fire sensitive and is generally killed even by low-intensity fires. Postfire reestablishment is via wind-dispersed seeds which readily germinate on fire-prepared seedbeds. The occasional mature tree which survives fire, those escaping fire in small, unburned pockets, and trees adjacent to burned areas provide seeds to colonize burned sites. Large trees occasionally survive light fires [31]. Scattered individuals or pockets of Engelmann spruce trees commonly escape burning because they occur in wet locations where fire spread is hampered. In subalpine habitats, scattered Engelmann spruce trees often escape fire because of discontinuous fuels, broken and rocky terrain, and the moist and cool environment [67,88]. Fire regime: Engelmann spruce-subalpine fir forests usually develop in cool, moist locations and experience fire-free intervals averaging 150 years or more [8]. Many Engelmann spruce stands are even aged, suggesting that they developed after fire [54]. Fuels and fire behavior: The fuel structure in stands dominated by Engelmann spruce and subalpine fir promotes highly destructive stand-destroying fires. Fuel loads are higher than in lower elevation montane stands, and the fuel beds tend to be irregular and have large amounts of needle litter accumulating under the narrow crowned trees [31,91]. The needles are small and fine, and form a compact fuel bed in which fire spreads slowly [28]. These concentrated, slow-burning fuels commonly produce flames high enough to reach Engelmann spruce's low-growing, lichen-draped branches and start crown fires [20,91]. POSTFIRE REGENERATION STRATEGY : crown-stored residual colonizer; short-viability seed in on-site cones off-site colonizer; seed carried by wind; postfire years 1 and 2 secondary colonizer; off-site seed carried to site after year 2

FIRE EFFECTS

SPECIES: Picea engelmannii
IMMEDIATE FIRE EFFECT ON PLANT : Engelmann spruce is easily killed by fire. It is very susceptible to fire because it has (1) thin bark that provides little insulation for the cambium, (2) a moderate amount of resin in the bark which ignites readily, (3) shallow roots which are susceptible to soil heating, (4) low-growing branches, (5) a tendency to grow in dense stands, (6) moderately flammable foliage, and (7) heavy lichen growth [87]. Crown fires typically kill Engelmann spruce trees. Engelmann spruce is also very susceptible to surface fires because fine fuels which are often concentrated under mature trees burn slowly and girdle the thin-barked bole or char the shallow roots [20,31]. Some large Engelmann spruce may survive light, surface fires, but these often die later due to infection by wood-rotting fungi that enter through fire scars [31]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Following fire, Engelmann spruce reestablishes via seeds dispersed by wind from trees surviving in protected pockets or from trees adjacent to burned areas. The rate of reestablishment is variable and depends on the proximity of surviving cone-producing trees and seed production during the year of the fire and immediate postfire years. In general, Engelmann spruce seedling establishment is very slow in areas burned by large, continuous crown fires because much of the seed source is destroyed. However, on small burns or near pockets of surviving trees within a large burn, Engelmann spruce usually establishes numerous seedlings within 5 to 10 years [42,44]. In areas where Engelmann spruce is abundant and lodgepole pine scarce before burning, Engelmann spruce establishes rapidly after fire if sufficient numbers of seed trees survive or are near the burn. If lodgepole pine is present in the preburn community, it usually seeds in aggressively, assuming a dominant role as it overtops any spruce seedlings establishing on the site [24,28,42]. However, Engelmann spruce seedlings usually survive under the developing pine canopy because of its shade tolerance. Above 9,850 feet (3,000 m), lodgepole pine does not regenerate, and burned areas remain open for several decades or longer. Postfire succession in this harsh, high-elevation zone (9,850 to 10,850 feet [3,000-3,300 m]) proceeds very slowly. Spruce slowly becomes established as scattered seedlings [12]. It may take 100 to 200 years before young spruce-fir forest covers the area. However, conditions in the upper parts of this zone sometimes make it difficult for tree seedlings to establish and survive at all. Here, grasses and sedges may form a mat which prevents tree seeds from reaching mineral soil [85]. Burned fir-spruce forest is replaced by alpine tundra which can persist for long periods of time [12]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : Postfire Engelmann spruce seedling establishment is best on moist surfaces where fire has consumed most or all of the duff leaving bare mineral soil. Seedlings do require some shade to survive; thus regeneration after fire is best on sites where standing dead trees, logs, or developing vegetation is present [73]. Engelmann spruce postfire regeneration is poor on sites subjected to high light intensities. A 26,000 acre (64,200 ha) burn on a high-elevation site in southwestern Colorado showed poor conifer regeneration 100 years after the fire. This was attributed to intense solar radiation which inhibited photosynthesis, causing a high percentage of spruce seedlings to die [75]. Postfire spruce regeneration is also poor where shrub and herbaceous cover is dense, where exposed mineral soil is subject to excessive evaporation, and where fire has only charred the duff [10]. Ash does not affect germination, but if it is deep, it can prevent a seedling's roots from reaching mineral soil [62]. In northern Colorado, 3 years after a late August wildfire in a dense, mature stand composed of Engelmann spruce, subalpine fir, and lodgepole pine, Engelmann spruce established 1,000 seedlings per acre (2,470/ha) in burned areas that were than less than 0.1 acre (0.05 ha). However, in the middle of the main burn, no Engelmann spruce seedlings had established by 3 years after the fire [10]. In Colorado, Peet [66] reported a 75-year-old burn that had good spruce regeneration near the burn boundary, but only 218 yards (200 m) inside the burn edge, few seedlings had established, and the area was still fairly open. Day [24] sampled lodgepole pine-Engelmann spruce x white spruce hybrid stands in southern Alberta that had established after fires that had occurred 29 and 56 years prior to sampling. He found that both pine and spruce had initiated large numbers of seedlings immediately after the fire. Pine, however, had established more seedlings and rapidly outgrew the spruce, forming a canopy that was three to four times taller than the spruce canopy. Pine seedling establishment had ceased by 30 years after the fire, but spruce continued to establish seedlings. Engelmann spruce eventually dominates sites where spruce and pine come in together after fire. The Research Project Summary Revegetation in a subalpine fir forest after logging and fire in central British Columbia provides information on prescribed fire and postfire response of plant community species, including Engelmann spruce, that was not available when this species review was originally written. FIRE MANAGEMENT CONSIDERATIONS : After clearcutting Engelmann spruce stands, broadcast burning can be used to prepare seedbeds for natural regeneration.  Broadcast burns which remove most of the duff or organic matter and burn hot enough to destroy some or all of the competing vegetation favor spruce seedling establishment [72].  However, seedling establishment is poor or nonexistent in areas where hot fires leave deep layers of ash or generate such intense heat that rocks are fractured, such as under slash pile fires [72,94].  For this reason, where large amounts of slash must be burned, windrows or piles should be kept small and cover a minimal portion of the area [3].  Engelmann spruce often occurs in cool and moist locations which restricts the time of year when effective broadcast burning can take place.  Prior to burning, duff must be dry enough to ensure that it will be consumed.  Seedling establishment will be inhibited on burns that only blacken the organic matter.  Some cull logs and slash should be left in place to provide shade and protection for developing seedlings [72]. Engelmann spruce stocking was greater than 50 percent and averaged 573 seedlings per acre (1415/ha) 5 years after broadcast burning in clearcuts in northern Idaho where the uncut stand composition was 56 percent western larch, 22 percent Engelmann spruce, 15 percent mountain hemlock, and 7 percent subalpine fir.  This broadcast burn exposed mineral soil on 53 percent of the area [14].  In northwestern Montana, Engelmann spruce seedling establishment was much greater on broadcast burned clearcuts where burning exposed mineral soil than on unburned clearcuts.  Eleven years after burning, stocking of Engelmann spruce seedlings was 23 percent on burned cuts but only 1 percent on unburned cuts.  Seventeen years after burning, stocking was 56 percent on burned cuts but only 2 percent on unburned cuts [84]. Broadcast burning is generally not recommended following partial cutting because residual Engelmann spruce trees are very fire sensitive.

REFERENCES

SPECIES: Picea engelmannii
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