Index of Species Information

SPECIES:  Populus grandidentata


Introductory

SPECIES: Populus grandidentata
AUTHORSHIP AND CITATION : Carey, Jennifer H. 1994. Populus grandidentata. 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 : POPGRA SYNONYMS : NO-ENTRY SCS PLANT CODE : POGR4 COMMON NAMES : bigtooth aspen largetooth aspen aspen TAXONOMY : The currently accepted scientific name for bigtooth aspen is Populus grandidentata Michx. (Salicaceae) [29,30]. Bigtooth aspen, along with five other aspen species, has been assigned to the subsection Trepidae of the section Leuce in the genus Populus. Because of their similarities, these six species are sometimes considered a single super species [44]. Bigtooth aspen and quaking aspen (P. tremuloides) are the only two North American aspen species. In literature concerning areas where both North American aspen species occur, many authors do not distinguish between bigtooth aspen and quaking aspen. The information is reported about "aspen" in general. In this writeup, "aspen" is used when citing studies in which both species are discussed collectively. Bigtooth aspen naturally hybridizes with the following species [29,30]: x P. tremuloides: P. xsmithii Boivin x P. alba (white poplar): P. xrouleauiana Boivin LIFE FORM : Tree FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY


DISTRIBUTION AND OCCURRENCE

SPECIES: Populus grandidentata
GENERAL DISTRIBUTION : Bigtooth aspen primarily occurs in the northeastern United States, southeastern Canada, and the Great Lakes Region. Its range extends from Virginia north to Maine and Cape Breton Island, Nova Scotia; west to southeastern Manitoba and Minnesota; south through Iowa to extreme northeastern Missouri; and east through Illinois, Indiana, Ohio, and West Virginia. Disjunct populations are found in Kentucky, Tennessee, North Carolina, and South Carolina [29,30]. ECOSYSTEMS : FRES10 White - red - jack pine FRES11 Spruce - fir FRES15 Oak - hickory FRES18 Maple - beech - birch FRES19 Aspen - birch STATES : CT DE IL IN IA KY ME MD MA MI MN MO NH NJ NY NC OH PA RI SC TN VT VA WV WI MB NB NS ON PE PQ BLM PHYSIOGRAPHIC REGIONS : NO-ENTRY KUCHLER PLANT ASSOCIATIONS : K095 Great Lakes pine forest K096 Northeastern spruce - fir forest K103 Mixed mesophytic forest K104 Appalachian oak forest K106 Northern hardwoods K107 Northern hardwoods - fir forest K108 Northern hardwoods - spruce forest SAF COVER TYPES : 1 Jack pine 5 Balsam fir 14 Northern pin oak 15 Red pine 16 Aspen 17 Pin cherry 18 Paper birch 19 Gray birch - red maple 21 Eastern white pine 25 Sugar maple - beech - yellow birch 32 Red spruce 33 Red spruce - balsam fir 35 Paper birch - red spruce - balsam fir 37 Northern white-cedar 43 Bear oak 46 Eastern redcedar 55 Northern red oak 60 Beech - sugar maple 108 Red maple SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Bigtooth aspen usually grows in even-aged mixed stands, most commonly with quaking aspen [29,43]. It is a codominant tree in both hardwood and conifer forests [13]. Bigtooth aspen does not occur as a subdominant species because of its extreme shade intolerance [38]. Quaking aspen is the predominant species in aspen stands in the Northeast and Great Lakes Region, but bigtooth aspen dominates on the drier upland sites [13,17]. Aspen stands dominated by bigtooth aspen are generally more open than those dominated by quaking aspen [17]. Overstory associates not previously mentioned in DISTRIBUTION AND OCCURRENCE include balsam poplar (Populus balsamifera), bur oak (Quercus macrocarpa), white oak (Q. alba), basswood (Tilia americana), black cherry (Prunus serotina), and sassafras (Sassafras albidum) [29]. A tall shrub layer is an important component of aspen forests [35]. Shrub associates include chokecherry (Prunus virginiana), downy serviceberry (Amelanchier arborea), dogwood (Cornus spp.), willow (Salix spp.), beaked hazel (Corylus cornuta), speckled alder (Alnus rugosa), American hazel (Corylus americana), and sweetfern (Comptonia peregrina) [13,29]. Bracken fern (Pteridium aquilinum) and dwarf bush-honeysuckle (Diervilla lonicera) are frequent subdominant understory species in bigtooth aspen stands [17,29,56]. Bigtooth aspen is listed as a dominant or codominant species in the following publications: 1. Wilderness Ecology: virgin plant communities of the Boundary Waters Canoe Area [35] 2. Aspen association in northern lower Michigan [17]

MANAGEMENT CONSIDERATIONS

SPECIES: Populus grandidentata
WOOD PRODUCTS VALUE : Bigtooth aspen wood is light colored, straight grained, finely textured, and soft. It is primarily used for pulp, but is also used to make particle board and structural panels. Minor uses include log homes, pallets, boxes, match splints, chopsticks, hockey stick components, and ladders [33,43]. Bigtooth aspen bark is pelletized for fuel and supplemental cattle feed [29]. IMPORTANCE TO LIVESTOCK AND WILDLIFE : Bigtooth aspen provides food and cover for wildlife. Moose and white-tailed deer browse bigtooth aspen [3,60]. Beaver eat bark, leaves, twigs, and branches [29]. Aspen provides the basic habitat for ruffed grouse over much of its range. Ruffed grouse feed on the leaves in the summer, staminate flower buds in the winter, and catkins prior to the breeding season. In feeding, ruffed grouse prefer quaking aspen to bigtooth aspen [20]. Approximately 116 nongame bird species breed in aspen-birch (Betula spp.) forests [5]. Cavity nesters use bigtooth aspen [21,65]. In a mixed hardwood forest in central New York, bigtooth aspen accounted for 25 percent of the trees with cavities although it made up only 12 percent of the potential cavity trees sampled [65]. PALATABILITY : Bigtooth aspen leaves are highly palatable to ruffed grouse [20]. NUTRITIONAL VALUE : Dry-weight bigtooth aspen browse averages 5.0 percent protein, 3.4 percent ether extract, 14.8 percent crude fiber, and 26.6 percent nitrogen-free extract [61]. The simulation of animal use by clipping did not alter the nutritional quality of bigtooth aspen browse [11]. Other normal foliar nutrient levels have been reported [9]. COVER VALUE : Bigtooth aspen provides cover for ruffed grouse. The best cover is provided by 5- to 25-year-old sapling stands with 3,000 to 8,000 stems per acre (7,000-20,000 stems/ha) [20]. VALUE FOR REHABILITATION OF DISTURBED SITES : Bigtooth aspen is of limited importance for revegetating coal mine spoils. Bigtooth aspen has been planted on mine spoils in Ohio and West Virginia [63]. Bigtooth aspen naturally regenerated on acidic sites in Pennsylvania, especially on sites where the soil had been ameliorated [26]. Bigtooth aspen regenerated naturally on barren, acidic, metal-contaminated soil near Sudbury, Ontario. The soil, contaminated by smelter fallout, had been treated with a surface application of limestone. Bigtooth aspen apparently colonized the site from off-site seed [67]. OTHER USES AND VALUES : NO-ENTRY OTHER MANAGEMENT CONSIDERATIONS : In the literature, the management of bigtooth aspen is rarely distinguished from that of quaking aspen. In order to regenerate a well-stocked vigorous aspen stand, the overstory must be removed [29,41]. Removing apical dominance stimulates aspen suckering [41,64]. If the parent stand is not harvested, it can be removed by shearing, chainsaw felling, girdling, treating with herbicide, or prescribed burning [41,42]. Discing and roller chipping are not recommended because these techniques damage the roots [41]. For optimum sucker density the parent aspen stand must have had at least 20 square feet basal area per acre (4.6 sq m/ha). At age two, an adequately stocked sucker stand exceeds 4,000 to 5,000 stems per acre (10,000-12,000 stems/ha) [41]. Although aspen groves thin naturally, additional thinning improves growth [29]. When thinning, bigtooth aspen should be favored over quaking aspen because of its superior growth and resistance to disease and insects, especially on dry sites [2,41]. Clones with superior growth and stem form should also be favored. Yields differ as much as 200 percent between clones on the same site [41]. Barnes [7] has developed a list of characteristics to use when distinguishing between clones in any given season. Only one thinning is recommended for aspen forests managed for for pulp. The stand should be thinned at age 30, leaving 240 trees per acre (590 trees/ha). Two thinnings are recommended if sawtimber is desired, one at age 10 leaving 550 trees per acre (1,360 trees/ha) and one at age 30 leaving 200 trees per acre (490 trees/ha). The final cut should be delayed for as long as the stand is healthy [41]. Repeated harvest of aspen on a site, especially by the whole-tree method, may reduce long-term productivity [2]. Short-term rotation management also reduces a stand's long-term health. Repeated short-rotation aspen harvest increases the colonization rate of Armillaria root rot (Armillaria mellea) [57]. Regeneration failure of aspen occurs if roots have been damaged by harvesting equipment. Aspen harvested on flat or gently rolling sites early in the summer may also have regeneration problems because of saturated soils [8]. Genetic improvement research on aspen has been conducted in the past two decades because of the increasing importance of aspen as a source of raw material for the pulp and paper industries. However, most recent work has concentrated on quaking aspen [4]. Bigtooth aspen responded to fertilizer (nitrogen, phosphorus, and lime in various combinations) with increased height, diameter, and volume growth during the 10-year period following the treatments. However, mortality of quaking aspen fertilized with nitrogen suggests further study is necessary before utilizing widespread fertilizer application in bigtooth aspen stands [49]. Herbicides control bigtooth aspen [34,66]. Bigtooth aspen is more disease resistant than quaking aspen. The most serious disease of bigtooth aspen is Hypoxylon canker (Hypoxylon mammatum) [29,68]. Other rots, fungi, and root decay affect this species [29]. Bigtooth aspen is a preferred host of gypsy moth. Death occurs when nearly complete defoliation by gypsy moth is followed by a fungal infection by Armillaria spp. [22]. By cutting trees and damming waterways, beaver destroy large aspen stands [29]. Biomass estimates have been reported for bigtooth aspen [36,39].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Populus grandidentata
GENERAL BOTANICAL CHARACTERISTICS : Bigtooth aspen is a native, short-lived, dioecious, medium-sized deciduous tree with a straight trunk and gently ascending branches. It is distinguished from quaking aspen by slightly larger leaves and large irregular teeth on the leaf edges. The fruit is a two-valved capsule [12,43]. Bigtooth aspen is a rapidly growing tree. At maturity, it attains heights of 60 to 80 feet (18-24 m) and diameters at breast height of 8 to 10 inches (20-25 cm). Stands begin to deteriorate after 50 to 70 years on good sites, but individuals may live as long as 100 years [29]. Bigtooth aspen is a clonal species. Clones resemble small groves consisting of many individual stems [7]. The roots of bigtooth aspen are shallow and wide spreading; the lateral root spread of a tree in a forest may be 33 to 66 feet (10-20 m). Generally, four to five lateral roots originate from the tree and then branch within 2 feet (0.6 m). Vertical, penetrating roots near the base anchor the tree [29]. The bark of young trees is smooth, but after three or more decades, it becomes rough and develops grooves [7]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : Bigtooth aspen regenerates by seed and vegetative reproduction. Bigtooth aspen is a prolific seed producer; a single tree may produce more than 1.5 million seeds [29]. Bigtooth aspen generally has good or better crops (greater than 61% of a full crop) 2 out of every 3 years [18]. The light seeds are dispersed long distances by wind [29]. Germination rates are high [14,29]. Seeds germinate under a wide range of temperatures as long as there is sufficient moisture. They will even germinate when submerged in water [14]. Despite high seed production and high germination rates, seedling establishment is uncommon. Few seedlings reach more than a few inches in height. Bare moist soil free of competition is necessary for seedling establishment. Short seed viability (2-3 weeks) also limits establishment. A seedling may grow 6 to 8 inches (15-20 cm) the first year [14,29]. Most bigtooth aspen forests regenerate vegetatively. When the parent tree is killed or the soil is heated, suckers develop from extensive, shallow lateral roots. Bigtooth aspen roots that produce suckers are generally less than 1 inch (2.5 cm) in diameter and about 3 to 7 inches (7.5-17.8 cm) deep. A sucker grows 3 to 6 feet (0.9-1.8 m) the first year, considerably more than a seedling. After a mature aspen stand is destroyed by fire or logging, roots may produce 3,200 to 24,000 suckers per acre (8,000-60,000/ha) [29]. Root suckers are initially dependent on the parent roots for water and nutrients. Their dependence decreases with time but is still substantial after 25 years. By age 25, the parent roots contribute the nutrient requirements for approximately half the yearly growth [68]. Multiple suckers result in a clone, a multistemmed vegetatively reproduced individual. Interclonal differences can be substantial, especially in sprouting ability [7,36]. Within a stand, clones are distinguished by sex, phenology, leaf morphology, disease resistance, bark and stem differences, branching habits, and other characteristics [7]. Sakai and Sharik [51] investigated the hypothesis that female bigtooth aspen clones would invest energy in fruit production at the expense of vegetative growth. They found no significant differences (P>0.05) in mean dbh or basal area density (basal area per unit clonal area) between male and female clones. The roots of bigtooth aspen may remain alive in a forest long after the last tree has died. The longevity of bigtooth aspen roots has not been documented. However, the roots of quaking aspen are known to persist in the absence of an aspen canopy. The roots are sustained by transient suckers that survive only a few years [54]. SITE CHARACTERISTICS : Bigtooth aspen most commonly occur on floodplains, gently rolling terrain, and the lower slopes of uplands. Large stands grow on sands, loamy sands, and light sandy loams. However, minor amounts of bigtooth aspen are found growing on almost any type of soil [29,41]. Bigtooth aspen has a lower soil pH limit of 4.0 [63]. Bigtooth aspen tolerates drier conditions than quaking aspen [2,17,47]. It has been classified as a subxeric species [10]. However, for good growth on upland sites the watertable must be at least 2 feet (0.6 m), but not more than 5 feet (1.5 m), below the ground surface. The soil must be moist but well aerated for good growth [29]. Bigtooth aspen ranges in elevation from sea level to over 3,000 feet (915 m) in North Carolina [29]. SUCCESSIONAL STATUS : Facultative Seral Species Bigtooth aspen is very shade intolerant [6,29,38]. It is a pioneer species on disturbed sites [13]. It persists in seral communities until senescence. Because of the tenacity and lateral extensiveness of its roots, aspen is able to regenerate and dominate disturbed sites that only had a minor aspen component in the original stand [13,44]. Palik and Pregitzer [38] reported no evidence of past suppression nor release of bigtooth aspen in a mature aspen forest. They suggest that most suppressed bigtooth aspen stems die. The rapid height growth of bigtooth aspen suckers allows it to outcompete other sprouting species such as northern red oak (Quercus rubra) and red maple (Acer rubrum) on many sites. Another reason for bigtooth aspen's propensity to dominate a site after disturbance is the large amount of space its lateral roots occupy. Oaks (Quercus spp.) and maples (Acer spp.) are generally limited to stump and root crown sprouts [38]. In the absence of disturbance, bigtooth aspen is replaced by conifers and hardwoods. On dry sites aspen is replaced by red pine (Pinus resinosa), oak, and red maple; on intermediate sites by eastern white pine (Pinus strobus); and on mesic sites by northern hardwoods, spruce (Picea spp.), and fir (Abies spp.) [13,37,56]. In the Great Lakes Region at the turn of the century, many mature pine forests were logged and burned. Bigtooth aspen and quaking aspen frequently dominated the postdisturbance forests [17,27,37,50,56]. Without fire or other disturbance, these forests are being replaced by later successional, shade-tolerant species [37,50]. In a study of forest succession in northern Michigan, bigtooth aspen, which dominated the postfire forest, was replaced by red maple and eastern white pine within 53 years [53]. On fine till soils in central New Hampshire, early successional species including aspen dominate postdisturbance stands. These species are replaced by sugar maple (Acer saccharum) and American beech (Fagus grandifolia) [69]. In the absence of fire, aspen-birch forests in Maine are succeeded by spruce [70]. SEASONAL DEVELOPMENT : Flowering in bigtooth aspen occurs in April or May, depending on air temperature. The seeds mature in May or June. Seeds disperse before the leaves are fully expanded. Bigtooth aspen flowers, foliates, and disperses seeds about 1 to 3 weeks later than quaking aspen in the same location [1,14,29].

FIRE ECOLOGY

SPECIES: Populus grandidentata
FIRE ECOLOGY OR ADAPTATIONS : Although bigtooth aspen is easily top-killed by fire, extensive vegetative reproduction, prolific off-site seed production, and the tenacity and lateral extent of its roots enable bigtooth aspen to perpetuate after fire [44,56]. Removal of the overstory and heating of the soil stimulate the sprouting of aspen roots [43,44,48]. Fire also creates a suitable seedbed and reduces competition [56]. Aspen almost always retains or extends its range following fire [15]. The extensive aspen root system allows it to dominate the postfire forest, even if aspen was only a minor component of the prefire stand [13,44]. Aspen roots persist an undetermined length of time in the absence of canopy aspen, making it possible for aspen to regenerate in a stand in which aspen was not even represented in the prefire overstory [44]. Aspen-dominated forests do not readily burn, especially when young and healthy [15,27,44]. Slow burning, low-severity surface fires are typical [15,25,27,48]. Decadent aspen stands contain more fuel and are more likely to burn than younger stands [15,44]. An understory of conifer species increases the flammability of aspen stands [24]. However, aspen is generally incapable of supporting a severe fire [25]. Crown fires in the surrounding forest generally drop into surface and ground fuels when they enter aspen stands [15]. Fire every 150 years may be necessary to maintain aspen [56]. Presettlement fires in aspen stands probably occurred most often in the fall when fuels are dry and leaf litter is deep [31]. The presettlement fire interval of aspen-birch-fir forests in the Great Lakes Region is estimated to have been about 80 years for very large [greater than 10,000 acre (4,000 ha)] fires [24]. Aspen-birch forests may have burned at intervals of 50 years or less [23]. Fahey and Reiners [71] estimated that aspen-birch forests in Maine had a 100-year fire interval during the 50-year period from 1909 to 1959. However, this estimate may be high because fires were declining during this time period due to fire suppression. Heinselman [24] suggested that aspen parklands had fire intervals of 10 years for large [1,001 to 10,000 acre (400-4,000 ha)] fires. Although quaking aspen is the principal tree in aspen parklands [24], bigtooth aspen occurs in the prairie-forest edge in Minnesota [19]. Historically, bigtooth aspen frequently occurred in vegetational patterns that are associated with fire. For instance, bigtooth aspen occurred with oak and quaking aspen along the west side of the Big Woods in Minnesota. These fire-tolerant species served as a firebreak between the frequently burned prairie and the fire-sensitive climax forest of the Big Woods. Aspen served as a fire break because grassland fires normally did not travel far into aspen stands due to low flammability. However, fuel would build up in the aging aspen stands and eventually an intense fire would destroy the aboveground biomass, rejuvenate the aspen, and eliminate the less fire-tolerant species. At the same time, frequent fire in the prairie prevented aspen expansion [19]. Another example of a fire-influenced vegetational pattern is in the Boundary Waters Canoe Area in Minnesota. Aspen commonly occurs in areas that frequently burn, such as large uplands areas distant from water and upwind of natural fire breaks such as lakes [23]. POSTFIRE REGENERATION STRATEGY : Tree with adventitious-bud root crown/soboliferous species root sucker Initial-offsite colonizer (off-site, initial community)

FIRE EFFECTS

SPECIES: Populus grandidentata
IMMEDIATE FIRE EFFECT ON PLANT : Bigtooth aspen trees are very susceptible to fire, although bigtooth aspen roots are very fire resistant. Bigtooth aspen bark is thin and does not protect the cambium from heat damage [29]. If there is sufficient fuel in a young sapling stand for a fire to burn, the fire will kill the saplings [44]. An average scorch height of 0.6 feet (0.2 m) will kill most aspen stems smaller than 6 inches (15 cm) in dbh [55]. Since most fires in aspen are of low severity, mature trees do not always succumb to fire. However, basal wounds caused by low-severity fire serve as entry points for disease organisms [29,44]. Prolonged drought and large amounts of slash are required to raise the soil temperature high enough to kill the roots of aspen [25]. Bigtooth aspen roots are deeper in the soil than quaking aspen roots, making root damage from fire highly unlikely in bigtooth aspen [40]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : A fire-killed aspen stand regenerates from surviving roots [25,29,44], which are stimulated to produce suckers when apical dominance is removed [44,64]. Roots must have a high cytokinin to auxin ratio to initiate sucker growth. If a low-intensity fire girdles aspen such that the downward movement of auxin in the phloem is interrupted but the upward movement of cytokinin continues in the xylem, the tree may survive for several years. Since the cytokinin continues to move out of the roots, it does not accumulate, and suckers do not develop [64]. Roots are stimulated to sprout if the soil is heated [25,29]. The soil is heated not only by the fire, but also by blackening of the soil surface and removal of overstory and duff [48]. The density and growth of suckers is dependent in part on fire severity. A low-severity fire that does not kill all of the overstory does not result in suckers as dense or vigorous as those produced after a moderate-severity fire [25]. Fire also prepares a favorable seedbed for bigtooth aspen. Seeds from off-site sources may blow onto burned sites and establish if there is sufficient moisture and if the competition is not severe [14,56]. Most shrub species in the understory of aspen forests are able to sprout after fire [13], as are many hardwood associates [38]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : Prescribed fire is used as a management tool to regenerate aspen stands [41]. Moderate-severity fire which kills all remaining canopy stems and removes duff is recommended following harvest to maximize the number of suckers [25]. A low-severity fire does not always induce sufficiently dense and vigorous suckers to regenerate an aspen stand [25]. Soil nutrient concentrations increased after wildfire on both clearcut and wholetree harvested sites that had formerly supported adjacent northern red oak-bigtooth aspen forests [73]. A mature aspen stand that is clearcut will generally have enough slash for fire to kill any residual standing trees. Fuel loadings of at least 10 tons per acre (22 t/ha) of slash less than 3 inches (8 cm) in diameter are recommended [41]. Refer to Beyerhelm and Sando [72] for fuel loading estimation techniques for aspen-northern hardwood stands. The recommended weather conditions needed to burn a harvested aspen site are reported for those stands with less than 25 percent conifers, greater than 25 percent conifers, and with little slash [41]. In northern lower Michigan, Scheiner and others [52] studied the vegetational response to clearcutting followed 1 year later by burning. Prior to cutting, bigtooth aspen dominated the site with 153 stems per acre (378 stems/ha). One year postcut, but prefire, the vegetation was dominated by red maple, northern red oak, and paper birch (Betula papyrifera) sprouts. However, in postfire year 1, bigtooth aspen was again the most abundant species with 9,911 sucker stems per acre (24,481 stems/ha). Although declining in number, bigtooth aspen continued to dominate the site for the duration of the study (5 years). Weber [64] studied the response of a quaking aspen and bigtooth aspen stand to four different treatments: surface burning before and after spring leaf flush and clearcutting before and after leaf flush. Three years after treatment, the preflush cut had the greatest stem density, average height, and basal diameter. Both cut treatments had greater stem densities than the burn treatments. The preflush cut treatment probably had more stems than the postflush cut treatment because carbohydrate root reserves were not depleted. One reason for low rates of suckering after prescribed burning was that the fires did not completely kill the overstory. The existing trees survived one to two postfire growing seasons and continued to show apical dominance, thereby preventing sucker development. The postflush fire resulted in more suckers than the preflush fire, probably because it was a hotter fire and killed more overstory trees [64]. Simard and others [55] developed equations for using postfire observations to predict fire-caused injury to and mortality in aspen. The National Fire-Danger Rating System uses the total ash and silica-free ash content of fuels as one of its variables. Ash is a noncombustible constituent of organic material and reduces the combustion rate of fuel. In a study in Michigan, the total ash content of dead bigtooth aspen litter fuels was 3.7 percent in the fall, 5.4 percent in the spring, and 6.2 percent in early summer. The silica-free ash content was 2.0, 2.8, and 3.5 percent for fall, spring, and early summer, respectively [31]. Ambrosia beetle (Xyleborus saxesceni) attacks fire-damaged bigtooth aspen [58]. Because of its low flammability, bigtooth aspen has been recommended for use in fire breaks, especially on droughty, sandy sites [27].

REFERENCES

SPECIES: Populus grandidentata
REFERENCES : 1. Ahlgren, C. E. 1957. Phenological observations of nineteen native tree species in northeastern Minnesota. Ecology. 38(4): 622-628. [74] 2. Alban, David H.; Perala, Donald A.; Jurgensen, Martin F.; [and others]. 1991. Aspen ecosystem properties in the Upper Great Lakes. Res. Pap. NC-300. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 47 p. [18412] 3. Allen, Arthur W.; Jordan, Peter A.; Terrell, James W. 1987. Habitat suitability index models: moose, Lake Superior region. Biol. Rep. 82 (10.155). Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service. 47 p. [11710] 4. Anderson, Neil A.; French, David W.; Furnier, Glenn R.; [and others]. 1990. A summary of aspen genetic improvement research at the University of Minnesota. In: Adams, Roy D., ed. Aspen symposium '89: Proceedings of symposium; 1989 July 25-27; Duluth, MN. Gen. Tech. Rep. NC-140. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station: 231-235. [12834] 5. Back, Gary N. 1979. Avian communities and management guidelines of the aspen-birch forest. In: DeGraaf, Richard M.; Evans, Keith E., compilers. Management of north central and northeastern forests for nongame birds: Proceedings of the workshop; 1979 January 23-25; Minneapolis, MN. Gen. Tech. Rep. NC-51. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station: 67-79. [18079] 6. Baker, Frederick S. 1949. A revised tolerance table. Journal of Forestry. 47: 179-181. [20405] 7. Barnes, Burton V. 1969. Natural variation and delineation of clones of Populus tremuloides and P. grandidentata in northern lower Michigan. Silvae Genetica. 18: 130-142. [2887] 8. Bates, Peter C.; Blinn, Charles R.; Alm, Alvin A. 1990. A survey of the harvesting histories of some poorly regenerated aspen stands in northern Minnesota. In: Adams, Roy D., ed. Aspen symposium '89: Proceedings of symposium; 1989 July 25-27; Duluth, MN. Gen. Tech. Rep. NC-140. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station: 221-230. [12828] 9. Blinn, Charles R.; Buckner, Edward R. 1989. Normal foliar nutrient levels in North American forest trees: A summary. Station Bulletin 590-1989. St. Paul, MN: University of Minnesota, Minnesota Agricultural Experiment Station. 27 p. [15282] 10. Cahayla-Wynne, Richard; Glenn-Lewin, David C. 1978. The forest vegetation of the Driftless Area, northeast Iowa. American Midland Naturalist. 100(2): 307-319. [10385] 11. Campa, Henry, III; Haufler, Jonathan B.; Beyer, Dean E., Jr. 1992. Effects of simulated ungulate browsing on aspen characteristics and nutritional qualities. Journal of Wildlife Management. 56(1): 158-164. [19705] 12. Chapman, William K.; Bessette, Alan E. 1990. Trees and shrubs of the Adirondacks. 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