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Quinney Natural Resources Research Library, The Bark Beetles, Fuels, and Fire Bibliography S.J. and Jessie E.

1973

The Ecological Role of Fire in the Area, Northwestern

Lloyd L. Loope

George E. Gruell

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Recommended Citation Loope, L. and Gruell, G. (1973). Ecological role of fire in the Jackson Hole Area, northwestern Wyoming. Quaternary Research, 3(3): 425-443.

This Article is brought to you for free and open access by the Quinney Natural Resources Research Library, S.J. and Jessie E. at DigitalCommons@USU. It has been accepted for inclusion in The Bark Beetles, Fuels, and Fire Bibliography by an authorized administrator of DigitalCommons@USU. For more information, please contact [email protected]. QUATERNARY RESEARCH 3, 425443 (1973)

The Ecological Role of Fire in the Jackson Hole Area, Northwestern Wyoming

LLOYD L. LOOPER AND GEORGE E. GRUELL~

Received March 20, 1973

Fire-history investigations in the Jackson Hole area of northwestern Wyoming reveal that most current stands of aspen and lodgepole pine regenerated following extensive fires between 1840 and 1890 and that widespread fires occurred in the 1600s and 1700s. White man’s major effect on the fire incidence has been the successful suppression during the past 39-80 yr. Successional changes in the absence of fire include the deterioration of aspen stands, massive invasions of subalpine fir in lodgepole pine stands, great increase in conifer cover, heavy fuel buildups in lodgepole pine and Douglas fir stands, and increase in sagebrush and other shrubs. Steps are being taken, starting in 1972, to allow fire to play a more natural role in and Yellowstone National Parks. Teton National Forest plans experimental prescribed burning to determine whether fire can stimulate successful aspen regeneration in the presence of large numbers of wintering .

T. S. Brandegee surveyed forest condi- due to successful fire suppression. Plant tions on the Teton Forest Reserve for the succession in the absence of fire has re- U.S. Geological Survey in 1897. His report sulted in an increase in the extent and aver- (Brandegee, 1899) stated : age age of coniferous forest stands and a deterioration of aspen stands. Shrub cover, It is only occasionally tkat tracts of timber especially big sagebrush, has increased of merchantable size are found, and areas markedly. Conifers continue to invade open containing notable quantities of merchant- able forest are few and limited. areas. (See Figs. 1, 2.) This condition appears to be due simply A primary management goal of the Na- and solely to fires which have swept over tional Park Service for Grand Teton Na- the country so completely ,and persistently tional Park is to maintain the park’s eco- that scarcely any part has been entirely ex- systems in as natural a state as possible empt from them, while nearly all portions have been burned again and again within a (Houston, 1971). The U.S. Forest Service generation . . . Under present conditions the has similar objectives for the Teton Wilder- tree-bearing regions as a whole decrease, while ness Area of the Teton National Forest. the aspen areas increase at the expense of The remainder of the Teton Forest is man- those now producing conifers. aged for multiple use, which involves com- The area of Teton Forest Reserve which mercial logging and management for wild- Brandegee surveyed now comprises most of life habitat, grazing of domestic livestock, the Teton National Forest and Grand watershed quality, and recreation values. It has become obvious to both land-man- Teton National Park. Ecological conditions agement agencies that an understanding of have changed over the past 75 yr, mainly the nature of pristine ecosystems and the 1 , Moose, Wyoming 83012. processesthat shaped them is necessary for ‘U.S. Forest Service, Jackson, Wyoming 83001. enlightened management.

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Copyright @I 1973 by University of’(Washington. All right.8 of reproduction in any form reserved. 426 LOOPE AND QRUELL

FIG. 1. Lower Gros Ventre Valley, Teton National Forest (Above). 1899. The landscape 20 yr after a major fire. Note the young aspen stands. The vegetation in the foreground is predominantly an herbaceous cover with scattered serviceberry and snowberry shrubs. Courtesy of Wyoming State Archives and Historical Department, Stimson Photo Collection. (Below). July 28, 1971. Aspen stands are mature and approaching decadence, 92 yr after fire. Aspen on distant bench has been largely replaced by Douglas fir. The foreground vegetation is now dominated by big sagebrush.

Our studies attempt to document the na- Forest and Grand Teton National Park, ture of the pristine vegetation patterns of with supplemental observations in Yellow- this area and the influence of fire on these stone National Park and Bridger National patterns. We try to determine the effects Forest. of fire and various stages of plant succes- sion on the outstanding wildlife populations THE AREA of the Jackson Hole area. Field investiga- Northwestern Wyoming has five major tions are mainly within Teton National ranges-the Teton, Absaroka, ECOLOGICAL ROLE OF FIRE, NORTHWESTERS WYO:\II’iG 127

FIG. 2. Lower Gros Ventre Valley (Above). 1899. Taken in the opposite direction from Fig. 1. The foreground of this scene shows a vigorous young aspen stand, which sprouted following the 1879 fire. Young conifers are becoming established under fire-killed snags on the distant slopes at right. Courtesy of Wyoming State Archives and Historical Department, Stimson Photo Collection. (Below). July 28, 1971. Reproduction is occurring at the margin of the 92-yr-old aspen stand, but not under the canopy. The distant slopes at right are now forested by a dense stand of Douglas fir. The lake formed following a landslide in 1925. 428 LOOPE AND GRUELL

Beartooth, Gros Ventre, and Wind River vicinity of our study area, the Hoback Ranges. Numerous lesser mountain ranges Range, the Mt. Leidy Highlands, the and uplands are present, including, in the Pinyon Peak Highlands, and the Yellow-

FIG. 3. Map showing location of study area in relation to the major physiographic features of northwestern Wyoming. ECOLOGICAL ROLE OF FIRE, NORTHWESTERN WYOMING 41’9

stone Plateau. Figure 3 shows the location cies at intermediate elevations (1850-2400 of the main study area in relation to the m) , characteristically occurring in almost major physiographic features of north- pure, nearly even-aged stands of fire origin. western Wyoming. Subalpine fir (A&es lasiocarpa), the most The hydrographic divide of the Teton abundant tree of subalpine forests, is found Range comprises the west boundary of from 1850 m to timberline at about 3050 Grand Teton National Park. The eastern m. Throughout the upper part of this zone, portion of the park lies entirely within fir is mixed with Engelmann spruce (Yicen Jackson Hole-a flat-floored valley about engebmannii) . Spruce sometimes occurs iu 65 km long from north to south and 15-25 almost pure stands in moist, fire-protectccl km wide, drained by the southward-flowing areas such as are found in canyons at the . Our major study area within east base of the Tetons. Whitebark pint Teton National Forest has been east of (Pinus albicaulis) , the other important southern Jackson Hole in the valley of the component of the subalpine forest, is found Gros Ventre River, which rises in the Gros from about 2400 m to 3050 m, and often Ventre . grows in nearly pure stands just below The area has a complex geologic history, timberline, and a great variety of bedrock types are Douglas fir (Pseudotsuga menziesii) is a exposed. The is composed pri- common species of the lower forest border marily of Precambrian granitic rocks. in this area. Scattered individuals occur up Paleozoic limestone and sandstone predom- to 2600 m on south-facing slopes of the inate in the Gros Ventre Mountains. Meso- Teton Range. Douglas fir forms extensive zoic shale and sandstone outcrop through- nearly pure stands on calcareous substrates out the Gros Ventre Valley. Many of these in the Gros Ventre Mountains and else- Mesozoic formations weather into unstable where in northwestern Wyoming. Limber clay soils, which are subject to sliding pine (Pinus flex&), less common than (Bailey, 1971). The Yellowstone Plateau is other conifers, is found at fores&sagebrush composed of Tertiary basalt and rhyolite. ecotones and on very dry south-facing Pleistocene glaciation has had profound slopes. Aspen (Populus tremuloides is fairly effects on the terrain. The valley of Jackson abundant at the lower elevations of the Hole is floored with thick, course outwash Jackson Hole area, although not nearly so deposited by glacial rivers. Numerous abundant as it is in parts of Utah and moraine deposits occur throughout the area. Colorado. Aspen often occupies the conifer forest-sagebrush ecotone and stands arc VEGETATION frequently mixed with Douglas fir. Blue spruce (Picea pungens), narrowleaf The vegetation of the Jackson Hole area cottonwood (Populus ungustifoliu) , balsam consists of a mosaic of nonforested areas popular (Populus balsumiferu) , and black and different-aged forest stands, most of cottonwood (Populus trichocurpu) occur on which originated following fire. Species floodplains in Jackson Hole. composition of parts of the mosaic is deter- Extensive nonforested areas occur in thiv mined not only by microclimate and soils, region. Sagebrush (including Artemisia tri- but in large part by the time since the dentuta and Artemisiu urbuscda) , bitter- latest fire and by the previous fire history, brush (Purshia tridentutu) , and rabbit- The forest portion of the mosaic is com- brush (Chrysothumnus spp.) are important posed essentially of eight speciea of the shrubs, which are associated with numerous family Pinaceae and four species of the species of grasses. Willows (Sulix spp.) and genus Populus. Lodgepole pine (Pinus con- sedges (Curer spp.) dominate swampy torta) is often the most abundant tree spe- sites. 430 LOOPE AND GRUELL

CLIMATE ment as a National Park in 1872. The first Climatic conditions vary widely in Jack- permanent settlers arrived in 1884 and 1885 son Hole because of the great variation in (Hayden, 1963). topography. Mean annual precipitation FIRE CLIMATE AND RECENT ranges from over 125 cm on the Pitchstone FIRE HISTORY Plateau of southwestern Yellowstone Park to less than 35 cm in the Gros Ventre Potential burning conditions vary greatly Valley. from summer to summer. The period of Moran, Wyoming, at an elevation of highest potential is between July 15 and 2075 m in northern Jackson Hole has a September 30. August 1531 is the most mean January temperature of -12OC likely period for extreme conditions to oc- WOW, a mean July temperature of cur. Periods of high fire potential are usu- 14OC (5B°F), and a mean annual precipita- ally brief and are terminated by an tion of 53 cm. At least half the precipita- abrupt change to cooler weather and/or tion falls as snow from November to April. precipitation. The average snow depth at Moran on April Data from cross sections of fire-scarred 1 is 100 cm. Summers in Jackson Hole are trees and weather records since 1916 suggest short and cool. Snow cover disappears from that conditions conducive to extreme fires valley areas in April and May and from have occurred at infrequent intervals. The high areas as late as July or August. May potential for ignition of extensive fires by and June are typically moderately wet lightning storms occurs infrequently, for months. July is the driest month, averaging most lightning storms in the area are ac- 2.5 cm of precipitation at Moran. Most of companied by precipitation. Small light- the summer rainfall falls in scattered thun- ning-caused fires were apparently frequent derstorms. The prevailing wind direction is in the past. Very large fires were rare, but from the southwest. they did occur periodically when a combi- nation of extremely dry conditions, one or more ignition sources, and high winds coin- WHITE MAN’S OCCUPATION cided. Data from fire-scarred trees show OF THE AREA that such conditions occurred in Jackson , who passed through Jackson Hole in the years 1856 and 1879, well be- Hole during the winter of 1807-1808, is gen- fore the initiation of fire-suppression erally regarded as the first white man to efforts. Extensive crown fires swept large enter the area. Several parties of fur trap- areas during these years. pers were in the area between 1811 and Crown fires occur here only under the 1820, and from 1820 to 1840 there was con- most extreme conditions. Even with very siderable fur trapping activity in Jackson dry atmospheric and fuel conditions, fires Hole. This activity ceased abruptly about generally remain in light fuels on the forest 1840, however, as a result of the sudden de- floor unless fanned by strong winds. The cline in popularity of the beaver hat and frequent occurrence of fire-scarred trees in the overexploitation of beaver populations. existing forest stands provide evidence of For the 1840-1860 period, there is little or creeping ground fires in the past. Observa- no evidence that white men were present tions made during the 250-hectare Pacific in Jackson Hole (Hayden, 1963; Mattes, Creek fire of August 22-29, 1971, in Teton 1946, 1948). National Forest, confirm this hypothesis. Miners, explorers, and hunters were This fire burned in a mixed stand of spruce, present in Jackson Hole periodically during fir, and lodgepole pine at an elevation of the 1860s and 1870s. Exploration of the 2459 m under extremely dry conditions but Yellowstone country led to its establish- was not pushed by high winds. Its general ECOLOGICAL ROLE OF FIRE, NORTHWESTERN WYOMING 431

behavior was a steady and thorough con- Since 1941, Teton National Forest has sumption of the moderately heavy ground had no fires as large as 400 hectares. Yel- fuels, with intermittent spotting and igni- lowstone National Park had two such fire,<, tion of the crowns. Only in areas of excep- totaling 1400 hectares. These figures reflect tionally heavy fuels, where the heat of the increasing effectiveness in fire detection and fire generated substantial convection cur- early suppression in northwestern Wyo- rents, did a crown fire develop. The crown ming. Fire suppression has been highly suc- fire died down in lighter ground fuels. cessful during the past 30 yr despite in- Prior to fire-suppression efforts, it is creased fuel buildups. The rarity of extreme likely that fires during most years were few burning conditions during this period has and small. Under burning conditions less probably contributed significantly to this than extreme, fires would die down and circumstance. smoulder. Periods of warm weather and in- creased wind would revive the blaze. In dry FIRE HISTORY PRIOR TO 1900 years, such fires could burn with varying Evidence from stand age and from sec- intensity for several months, affecting large tions of fire-scarred trees has been used to acreages. investigate the fire history of Grand Teton Organized fire suppression dates back to National Park and parts of Teton National the establishment of Teton Forest Reserve Forest. Numerous very small fires have oc- in 1897. The diary of B. F. Bondurant, a curred in the Teton Range because of fre- fire guard employed by the U.S. Forest Ser- quent lightning strikes, but large fires have vice, describes how he extinguished a fire, been rare because of rugged topography started by a lightning strike, which burned and numerous natural firebreaks-espe- one-half acre in Hoback Basin on August cially bare rock. In contrast, no extensive 4, 1904 (Sargent, 1963). The U.S. Army area in the valley of Jackson Hole has cs- was extinguishing fires in parts of Yellow- caped fire for much over 200 yr, and most stone Park in the 1880s according to areas burned in the interval between 1850 Superintendents’ Reports (Harris, 1886; and 1885. Fire-scar and standage data sug- Boutelle, 1890). Detection and suppression gest that large fires occurred about 1765. of fires improved markedly in the 1920s in the early 1840s about 1856, and in the and 1930s with the establishment of fire interval of 1878-1885. Most forest stanfls lookouts throughout most of the forested in the valley originated following the fires area of northwest Wyoming. The remark- of 1856 and 1879. Several square kilomet,cra able success of suppression efforts is re- of lodgepole pine forest burned in the north- flected in the fact that a total area of about ern Teton Range as recently as 1910. 125 hectares has burned since 1910 in what Sampling from the Gros Ventre Valley is now 130,000-hectare Grand Teton Na- in Teton National Forest, contiguous with tional Park. Highly effective suppression of the eastern edge of Jackson Hole, indicat,es fires in inaccessible parts of Northwestern that fires burned somewhere in the drainage Wyoming did not begin until about 1940. during each decade of the 19th century. Moderate detail is avaiIable for fire However, fires seem to have burned extcn- records since 1931. In the 1931-1940 period, sive arcas only during the 184Os,187Os, and there were seven fires in Teton National 1880s. Most forest stands in the Gras Forest larger than 400 hectares each. The Ventre drainage were affected by one or total area burned by these fires was over more fires during the 1840-1890 period. 14,000 hectares. During this same decade, Many stands at higher elevations in the Yellowstone National Park had nine fires Gros Ventre Mountains burned during this over 400 hectares, totaling about 18,000 period, too, but other stands escaped. Many hectares. open lodgepole pine stands contain numcr- 432 LOOPE AND GRUELL ous individuals with two or three fire scars yr. Stands at high elevation or protected from this period. Some of the largest fires by topographic peculiarities are typically occurred about 1842, 1872, and 1879. influenced by fire less often. Tundra habi- Limited sampling of sections of 300-400- tats may be virtually free of fire’s influence. yr-old trees in the Jackson Hole area (mostly Douglas fir) with multiple fire SOURCES OF IGNITION scars suggest that numerous fires occurred There can be little doubt that forest fires in the 1600s and 1700s. Trees of this age have had a profound influence on the may have three or four fire scars, even ecosystems of northwestern Wyoming for though a fire generally scars only a small centuries. Organic material produced by percentage of surviving trees in its path, photosynthesis in regions with cool climate The best evidence for comparative fire accumulates at a much more rapid rate than frequencies in northwestern Wyoming in it decays, creating a fuel source (Beaufait, the 17th, l&h, and 19th centuries comes 1971). During very dry conditions of late from Houston (1973). Houston invest.i- summer or early fall, which are attained gated the fire history of the winter range in occasional years, adequate ignition can for ungulates of northern Yellowstone produce extensive conflagrations. The fire Park. Cross sections were obtained from 33 frequency at a given site partially depends live and 10 dead fire-scarred Douglas firs on the frequency of ignition. Two major located at forest-grassland ecotones in the sources of ignition exist-lightning and Lamar, Yellowstone, and Gardner River man. valleys. Vegetation of this area is com- Nowhere in the world is lightning such parable to Jackson Hole but somewhat a significant ignition source as in the conif- drier and warmer. Trees averaged 308 yr erous forests of western old, and eight were over 400 yr old. Individ- (A. Taylor, 1971). About 60% of the mod- ual sampled sections showed an average ern forest fires in this area are caused by frequency of scarring of about 50 yr, prior lightning. Data from northwestern Wyo- to white man’s influence in reducing fire ming support this assertion. Lightning ig- frequency. Comparison of scars from nited 53% of 857 fires from 1931 to 1971 adjacent trees suggests one fire per 20-30 in Teton National Forest. In Yellowstone yr. The occurrence of similar fire dates over National Park, 56% of 1228 fires between wide areas suggests the likelihood of eight 1931 and 1967 were ignited by lightning or ten large fires in the past 300-400 yr. (D. Taylor, 1969). That lightning starts Fire frequencies were very similar during fires now and has caused them in the past is the 17th, l&h, and 19th centuries. The most undeniable. We have good data on the recent fire scars were generally no later relative importance of lightning vs man as than the 1870s. Only two of the sampled a source of ignition at present. How did the trees on the major study units were scarred past compare with the present in this in the 1890s and none of the sampled regard? Douglas firs had been scarred since 1900. It is possible only to speculate on the im- A comparison of Houston’s data with our portance of aboriginal populations in influ- data from Jackson Hole suggests that fire encing the past fire frequency in the Jack- occurred less frequently in Jackson Hole son Hole area. Artifacts from the north end than in the grassland areas of northern Yel- of suggest seasonal occupany lowstone Park. Few 400-yr-old Douglas firs in this valley over 10,000 y.a. (Frison, in our area exhibit more than four fire 1971). More concrete evidence exists for the scars. For most low-elevation sites in Jack- periodic occupation of Mummy Cave, a site son Hole, we estimate the mean natural fire 105 km northeast of Jackson Lake, for the frequency to have been one fire per 50-100 past 9000 yr (Wedel et al., 1968). Abun- ECOLOGICAL ROLE OF FIRE, NORTHWESTERN WYOMING 433 dant evidence points to the presence of man Yellowstone National Park, prospecting in Jackson Hole during the past 4500 yr. and hunting trips, and initial white settlc- groups are believed to have ment of Jackson Hole. Travel and habita- hunted bison and antelope there (Frison, tion in the area were still infrequent, how- 1971). ever (Doane, 1877; Baillie-Grohman, 1884; The use of fire by aboriginal man in vari- Buxton, 1893; Fryxell, 1932). The remote ous parts of the world to drive game in setting, harsh environment, and limited hunting is moderately well documented mineral wealth of Jackson Hole deterrecl (Stewart, 1956). Stewart considers unextin- occupancy. Permanent settlement did not guished campfires to have been a major ig- begin until 1884. Native accounts com- nition source for fires in prehistoric times. pletely lack instances of wildfires caused by No specific information is available for the white man or Indians during this periocl. *Jackson Hole area. Early man may have As previously stated, tree-ring evidence been a significant source of ignition there. indicates extensive fires during the The elimination of aboriginal man from the 1870-1885 period, especially about 1872 ant1 scene in the late 19th century eliminated 1879. Historic accounts document fires iI1 this potential ignition source. 1878,1879, and 1880. W. W. Jackson passed In many areas of the west, the explora- through Jackson Hole in the late summe tion and settlement by white man clearly of 1878 and photographed the Tetons from increased the fire incidence. Timbering, Signal Mountain. He was unsatisfied u>ith land clearing, railroads, and campfires were the quality of his photographs “because of constant sources of ignition, causing some a smoky haziness (caused by forest fires irt of the largest fires in recorded history the vicinity) that filled the air” (Jackson (Holbrook, 1943). Militant Indians were and Driggs, 1929). Thomas Moran visitctl blamed for many fires in the Rocky Moun- the west side of the Tetons in August 1879. tains during the late 19th century (e.g., Conditions for making sketches were far Hanna, 1934; Town, 1899), though some from ideal throughout the trip because accounts suggest that their incendiary role smoke from forest fires became “so dense has been overestimated by white historians as almost to obscure the peaks of the (Brown, 1971). Tetons” (Fryxell, 1932). W. A. Baillie- Most evidence suggests that the white Grohman camped at Jenny Lake in August, man did not greatly increase the fire fre- 1880 but was forced to leave the area when quency in Jackson Hole during the 19th chased by a forest fire several miles to the century. An occasional wildfire undoubt- north of *Jenny Lake that was “running he- edly resulted from a poorly placed or un- fore a northwesterly breeze” (Baillie-Groh- attended campfire during the 1820-1840 man, 1884). In these accounts, no mention furtrapping period. However, data from is made of fires being caused by humans. fire-scarred trees indicate that fires during Observers of forest conditions in north- this period were less frequent than during western Wyoming in the 1880s and 1890s most decades of the 1800s. There is little felt that white man’s carelessness was at, evidence of white man in the Jackson Hole least partially responsible for the devastn- area between 1840 and 1860 (Mattes, tion of forests by fire. Hague (18861 recog- 1948). Only a few records exist of his pres- nized that “evidence of forest fires 160 ence there in the 1860s (Raynolds, 1868; years ago may still be traced,” but felt that De Lacy, 1876). “disastrous fires took place, caused by t,he There is abundant documentation for the carelessness of white man” following es- presence of white man in the region during tablishment’ of Yellowstone Park. He the 1870s and early 188Os, with the advent felt that “with the coming of the rnil- of government surveys, establishment of road and the erect.ion of hotels, fires stead- 434 LOOPE AND GRUELL ily increased,” until strict precautions were marked reduction of fire frequency during enforced by the U.S. Army in the late the past 30-70 yr. 1880s. Brandegee’s (1899) comments for the Teton Forest Reserve are vague as to ROLE OF FIRE IN MAJOR the cause of the fires. Taylor (1969) points ECOSYSTEM TYPES out that the importance of lightning in ig- niting fires was not appreciated in the 19th Aspen century. He determined that Superinten- The role of fire in aspen ecology is of spe- dents’ Reports for Yellowstone National cial concern to land-management agencies Park prior to 1899 failed to mention light- of northwestern Wyoming because of the ning-caused fires. Fires were thus assumed current deteriorated condition of most to be man-caused. aspen stands. The failure of aspen repro- One cannot state with certainty that duction has been a concern in Jackson Hole white man did not significantly influence for over 25 yr. Traditionally, this has been the natural fire frequency in the Jackson blamed on “overbrowsing” by the Jackson Hole area during the period of exploration Hole elk herd, which was believed to exceed and settlement. Certain lines of evidence numbers that existed in primitive times. suggest this, however. These include: More recent evaluations of historic elk 1. Fire-scar evidence from 300 to 400- numbers suggest a decrease in numbers yr-old trees seems to indicate a fire fre- since primitive times (Cole, 1969; Gruell, quency during the 1600s and 1700s roughly 1973). Some changes in elk distribution similar to that during the 1800s. have occurred, particularly adjacent to 2. There is apparently no correlation be- winter feedgrounds, but in general the tween frequency of people in the area and forage utilization patterns resemble those frequency of conflagration. (Large areas of the past. Studies of fire history and fire burned in the 1840s and 185Os, when very ecology suggest that the decline of aspen few or no white men were in the area. Only is primarily due to an alteration of the small numbers were present during the natural fire regime. great fires of the 1870s and early 1880s. In the , aspen repro- After settlement began in the mid-1880s, duces almost entirely by sending up vegeta- few extensive fires occurred.) tive shoots (suckers) from the root system 3. Narrative accounts are almost entirely of a clone, and there is apparently very lit- lacking for accidentally or intentionally set tle reproduction by seed. Aspens are readily fires. Such documentation is abundant for killed by fire. Large numbers of vegetative other areas of the west. sprouts are produced following a fire in an In summary, it appears that the fire aspen stand ; 12,000-20,000 stems per hec- regime for the 19th century dedl,ced from tare is not unusual. Hague (1886) noted fire-history investigations in modern forest that in northern Yellowstone Park aspen stands can be interpreted as fairly typical is “the first tree to spring up upon recently of pristine conditions. Adequate ignition burned areas” and that “by so doing, it sources existed before the coming of white helps to conceal unsightly charred trunks, man to northwestern Wyoming. Aboriginal and adds bright color to the somber hill man probably started fires, intentionally or slopes.” inadvertently, but sufficient lightning fires Apparently, fire produces a physiological have been ignited by late-summer storms stimulation of the root system to send up to assure the existence of fire-influenced suckers, which is tied to the killing of ma- forests with periodic cyclic disturbances ture trees of the clone and is thought to be without human ignition. White man’s influenced by auxins (Farmer, 1962). Re- major effect on the fire incidence has been moval of competing vegetation and the ECOLOGICAL ROLE OF FIRE, KORTHWESTERiX WYOMING 435 cleansing of the stand by disease are other (now and in the past), aspen reproduction factors involved (Graham et al., 1963). Nol and establishment, though most prolific fol- only are large numbers of suckers produced lowing the fires of X450-1890, has continued following a fire, but these suckers have sporadically until the present. However, in rapid growth. most instances, reproduction during the Competition among suckers as the stand past 70 yr has been insufficient to rcplnccb develops causes rapid reduction in the num- the deteriorating stands. ber of stems per hectare. However, sucker- Individual aspens 200 yr old rarely W- ing continues for several years after a fire. cur. Stand deterioration usually sets in In the fall of 1971, we made a study of the after an age of about 80 yr is reached. relationship of tree ages in aspen stands Various diseases, the most important of of the Jackson Hole area to the date of the which are sooty bark canker and heart rot, most recent fire influencing each stand, are the immediate cause of the death of the which showed that aspen suckering clue to trees. Invasion of conifers (limber pinch, stimulation by fire continues for 10 yr or Douglas fir, lodgepole pine, subalpine fir, more after the fire. This may be largely due and Engelmann spruce) occurs in many iui- to the expansion of clones following fire. tially pure aspen stands and smctimt’s The age structure of present stands is re- contributes significantly to the clcolinc of lated to levels of browsing by elk and the stands. Mixed stands of aspen anri moose, which are determined by such fac- conifer originated following fire on son~e tors as population densities, forage prefer- sites. In most of these stands, the conifers ences, topography, and availability of have attained dominance and acc:clcr:~te~l forage species other than aspen. We have the death of aspens through shntling anI1 found no evidence to suggest drastic in- root competition. creases in elk browsing from natural con- Nearly all aspen stands in the .Jnckson ditions, except in the vicinity of sites where Hole area had their origins following fires elk are unnaturally concentrated when between 1850 and 1890. Very little repro- supplied supplementary artificial feed in duction has occurred since 1900. Most winter. There has been a great increase stands are now either in or approaching a in moose browsing in the past 50 yr. critical period. If no fires occur in the nest In areas where ungulate browsing is 20-50 yr, many clones will be reduced IO naturally heavy at present (and presum- a situation in which they no longer c(;~I- ably was heavy in the past), all aspen trees tribute significantly to the vegetative cover. that sprouted following the fires of the In areas of moderate to light game use, 1850-1890 period became established within reproduction of aspen on stand margins and 1030 yr after the most recent fire. The under deteriorating stands has been much greatest intensity of reproduction was gen- greater during the past 20 yr than during erally in the 5-yr period immediately fol- the previous 50 yr (1900-1950). We at- lowing the fire. Virtually no aspens have tribute this increase in reproductive vigor become established in these stands within to the deteriorat’ion of parent stands, which the past 70 yr. For example, in a stand accelerated about 20 y.a., when most aspens near Tent Creek in the Gros Ventre valley, reached an age of 70-80 yr. which originated following a fire 120 y.a., In summary, successful regeneration of all aspens sampled (19) were over 90 yr aspen stands in the Jackson Hole area was old. In a stand near the mouth of Goose- stimulated by fire in the past. Some sucker- wing Creek, which originated following a ing occurs now without fire, and, in areas fire 98 y.a., all aspens sampled (9) were at with light game use, such suckers can sur- least 89 yr old. vive to produce mature trees. Even here, In areas where ungulate browsing is light however, reproduction is generally not suffi- 436 LOOPE AND GRUELL cient to maintain the stand. In areas with millennia. The high percentage of lodgepole heavy game use, suckers rarely survive. stands in a susceptible stage at present, due to fire suppression, has presumably resulted Lodgepole Pine in high beetle populations over an unprece- Lodgepole pine is well known as a species dented expanse of forest. Photographs that thrives under the influence of fire taken before 1900 show that some mature throughout most of the Rocky Mountains. stands of lodgepole were decimated by In many parts of the Rockies, lodgepole has mountain pine beetle before the advent of predominantly serotinous cones, which re- successful fire suppression, however. main closed for many years unless heated Within IO-20 yr after a fire in a lodge- above 45OC (113OF) (Fowells, 1985). In pole stand, a dense level of stocking is usu- northwestern Wyoming, however, serotiny ally attained. Typical 90-yr-old stands is relatively unimportant. Most cones open sampled in Jackson Hole have 1700-2300 without heat, and the seeds are scattered trees per hectare larger than 10 cm d.b.h. without fire. However, lodgepole pine has Many dead stems of suppressed individuals abundant seed production and is a success- are usually present. The forest floor is ful invader after fire if there is an adequate highly shaded in such a stand, and there seed source nearby. The reproductive po- is generally no lodgepole reproduction. Re- tential of nonserotinous lodgepole is sug- productive success of subalpine fir in the gested by the establishment of large num- understory is variable. At an age of about bers of seedlings on mineral soil exposed in 120 yr, typical lodgepole stands have roadcuts. The widespread occurrence of liv- 80&1050 stems per hectare. Most stands of ing fire-scarred lodgepoles suggested that this age have been affected considerably by individual trees often survived past fires mountain pine beetle attack. With in- and served as a seed source for t.he new creased sunlight from thinning of the stand stand. The probability also exists that, and reduced competition, fir reproduction when a fire killed an entire stand, enough increases. seeds from unopened mature cones survived At an age of about 200 yr, only about the fire to contribute substantially to the 400-650 mature lodgepole pines per hectare seed source for a new stand. remain. The stands have suffered high mor- After lodgepole pine stands attain an age tality from mountain pine beetle and be- of 80-100 yr (and the trees a diameter of come increasingly susceptible to windthrow. 20-25 cm), they become susceptible to at- Large openings in the canopy occur, and tack by mountain pine beetles (Dendroc- lodgepole reproduction commences. In tonus ponderosae), which lay eggs in the many 200-yr-old stands, the number of phloem so that trees are girdled by the lodgepole seedlings and saplings is greater feeding larvae. As age and diameter in- than those of fir. In the absence of fire, such crease further, lodgepoles become increas- stands might take several hundred more ingly susceptible to attack (Roe and Am- years for fir to attain dominance. The sub- man, 1970). Most stands in the area have stantial fuel accumulation in old lodgepole thus been recently vulnerable to beetle at- pine stands predisposes such stands to fires, tack, which has killed 10-45s of the trees which perpetuate lodgepole dominance. over 5 in. (13 cm) d.b.h. (diameter at breast height) in much of Grand Teton Na- Subalpine Fir tional Park and Teton National Forest in Subalpine fir might be termed the “cli- the past 12 yr (Klein et al., 1972). The max” conifer species for much of the Jack- mountain pine beetle, a native insect, is a son Hole country. It has increased greatly highly significant ecological factor that has in importance due to fire suppression and influenced lodgepole pine ecosystems for is the primary species invading lodgepole ECOLOGICAL ROLE OF FIRE, NORTHWESTERN WYOMING 437 pine stands and many aspen stands. Its in- the east slope of the Canadian Rockies in creasing importance and its value as moose Alberta. browse may have been a contributing fac- Where an adequate seed source and suffi- tor to the increase in moose population cient moisture are present, spruce regener- levels in Jackson Hole since 1900. ates following fire and may form mixetl Fir, like spruce, is easily killed by fire. stands with lodgepole pine. Douglas If an abundant seed source exists near a fir often establishes individuals in such :t burn, fir can become established in a mixed postfire stand. Since few lodgepoles survive stand with lodgepole. A pure, dense lodge- beyond the age of 200 yr, an almost pure pole stand sometimes has little fir reproduc- stand of spruce occupies the site from 200 tion at an age of 120 yr. Generally, by an to 500 yr following a fire. During this stagth. age of 80-100 yr, substantial fir reproduc- subalpine fir often comprises less than 109, tion has begun in a lodgepole stand. With- of the basal area of the stand. But spruce out fire, shade-tolerant fir would virtually regeneration is minimal after the lodgepolc replace lodgepole in most stands within canopy closes. As the age of the stand in- 250-400 yr after a fire. Fir invasion, how- creases, spruce reproduction approaches ever, contributes to the fuel buildup. In zero, except on rotting logs and in openings combination with the fuel from dead lodge- with exposed bare mineral soil. The ac- pole pines, a fire that will reinitiate the cumulation of an organic layer at the soil cycle becomes likely. surface and the low light penetration apparently inhibit spruce seedling estab- Engelmann Spruce lishment. Seedlings of subalpine fir are Reed (1969) and Oosting and Reed lo-20 times as abundant as those of spruce (1952) found spruce-dominated forests at in these stands. Oosting and Reed (19521 elevations from 2700 m to timberline in the found a similar ratio of fir seedlings to Wind River and , spruce, but felt that high mortality of respectively. In the , young fir resulted in maintenance of a nearly pure spruce stands are largely con- climax spruce community. The nearly fined to the higher elevations (2400-3050 even-aged structure of the dominant species m). In the Teton Range, spruce-dominated here strongly suggests that this community forests are found primarily in sheltered is not “climax.” canyon bottoms and slopes from 2080 m to It appears that fir would eventually re- 2400 m. The spruce stands of the Tetons place spruce here in the prolonged absence are characteristically on the west sides of of fire. The tremendous fuel buildups that lakes at the base of steep slopes. Fires are accumulate as the spruce canopy breaks up therefore much less frequent than in less predispose the stands to fire. No present shehered areas. Large individuals of spruce, stands exhibit a late stage in the hypotheti- up to 1.2 m in diameter, occur in these cal succession from spruce to fir. forests, which are located mainly at the Spruce is conspicuously uncommon in east base of the Teton Range. lodgepole forests of valley areas of Grand Spruce trees are easily killed by fire, and Teton National Park. This appears to be regeneration is usually slow following fire. partly due to past fire history, which has The age structure of these stands suggests obliterated a seed source, and partly due that even these old-growth spruce forests to moderately dry late-summer conditions. depend on fire for their establishment, Fir seeds are larger than spruce seeds and because the stands consist of nearly even- contain much more food for utilization by aged spruce, with very little spruce repro- seedlings (Fowells, 1965 ; Oosting and duction. The situation resembles that de- Reed, 1952). Seedling fir can develop a root scribed by Bloomberg (1950) for spruce on system more rapidly than spruce-an im- 438 LOOPE AND GRUELL

portant adaptation where soil moisture is age structure on north-facing slopes of marginal. A few mature spruces occur in , a prominence in soutbeast- these valley lodgepole pine forests, and oc- ern Grand Teton National Park, illustrate casional seedling establishment occurs. particularly well the past role of fire and In the Gros Ventre Valley, spruce com- the need for the return of fire to natural monly invades lodgepole pine and aspen ecosystems. Lightning fires swept across the stands on north-facing slopes. In this area, sagebrush flats in the past and ignited these a seed source for spruce survived past fires forests every 25 to 100 yr. Large, thick- in moist pockets. Moisture from seepage barked Douglas firs, up to 1.6 m in diame- often contributes to spruce establishment. ter, have as many as four or five fire scars, some of which date back to the 1600s. The Whitebark Pine most recent fire, which burned in 1879, Above 2600 m, whitebark pine often affected most of the Butte and left only the colonizes areas in spite of its large seeds, larger trees. which require transport by squirrels, chip- From field observations and from reports munks, and birds for dispersal. Near in the literature for similar ecosystem types Amphitheatre Lake in the Teton Range a (Lyon, 1971), the course of plant succession nearly pure stand of whitebark pine has be- since the 1879 fire can be postulated. In the come established following a fire about 200 summer following the burn, a lush growth y.a. North of Mt. Berry, in the northern of herbaceous species such as fireweed be- part of the Teton Range, mixed stands of came established, due to an abundant whitebark pine and lodgepole pine occur on source of nutrients and full sunlight. an area that burned about 225 y.a. The Scouler’s willow and quaking aspen, sprout- forest on the summit of Mt. Berry burned ing from undamaged root systems, attained about 90 y.a. The reproduction of this heights of l-2 m during the first year and burned area is about 75% whitebark pine continued rapid growth during the next 20 and 25% lodgepole pine. yr. The invasion of Douglas fir seedlings was slow. Douglas Fir After about 50 yr, such a dense growth Extensive stands of Douglas fir occupy of Douglas fir saplings was established that the lower slopes of the Gros Ventre Moun- competition became intense. The trees grew tains on sedimentary rocks and north-fac- in such proximity that branches on the lower ing slopes at low elevations in eastern Jack- half of the trunks died. Aspens and willows son Hole. In much of northern Jackson died or persisted as relict individuals. The Hole, Douglas fir is found in speciaI dark forest floor was covered with shade- edaphic situations-shallow soils, ridges, tolerant pinegrass. and south-facing slopes. On higher-eleva- In 1972, 93 yr after the 1879 fire, there tion sites, Douglas fir may be seral to is a considerable fuel buildup on Blacktail subalpine fir, but it maintains itself for Butte from dead Douglas fir branches and several centuries and is perpetuated by wood skeletons of aspen and Scouler’s periodic disturbance. willow. If a fire were to reach these stands, Fire in the past played a major role in most of the young Douglas firs would be Douglas fir forests by maintaining an open killed, but many of the large, thick-barked condition through periodic ground fires. trees would probably survive. Crown fires undoubtedly occurred in areas If fire suppression is continued, it is pos- of heavy fuel accumulation. The thick bark sible that fire could be prevented from con- of Douglas fir allows mature trees to sur- suming the Blacktail Butte Douglas fir vive most ground fires. stands for another 50-100 yr, but not in- Pure stands of Douglas fir with a mixed definitely, In the interim, fuel accumula- ECOLOGICAL ROLE OF FIRE, NORTHWESTERN WYOMING 439 tions would continue to build up to the vasion of grasslands. Reduction of fine fuels point where a fire would very likely kill all due to livestock foraging has had a definite Douglas firs-even those with the thickest influence in reducing fire frequency bark. The ecological consequences of such throughout much of the west (Leopold, a fire would likely prove highly unfavor- 1924; Humphrey, 1958). able, since aspen and willow would have de- Fire greatly influences the shrub com- teriorated beyond the point of potentially position of various community types. successful vegetative reproduction. Also, Whereas big sagebrush and low sagebrush the seed source for Douglas fir would be are killed even by low-intensity fires, many drastically diminished. shrub species are stimulated by fires. Willows (S&-c spp.), silver sagebrush (Ar- Sagebrush and Other Shrubs temisia cana), bitterbrush (Purshia trident- Fire periodically swept the sagebrush ata), rabbitbrush (Chrysothamnus nause- flats of Jackson Hole and the Gros Ventre OSZLS), serviceberry (Amelanchier alnifolia) , Valley. Big sagebrush and low sagebrush chokecherry (Prunus virginiana var. (Artemisia trident& and Artemisia arbus- melanocarpa), snowberry (Symphoricarpns cula) are readily killed by fire. Grasses and SPP.), mount’ain ash (SorblLs SCOPU- other herbaceous plants flourish following l&a), and snowbrush ceanothus (Cenno- this nutrient release and reduction of com- thus velutinus) are some of the shrubs of petition for soil moisture. Sagebrush pro- northwestern Wyoming that may resprout duces abundant seeds and is capable of after aerial parts are killed by fire. Witch rapid reinvasion on sites where competition many of these species, the stage of growth, from grasses is less severe. The grass cover the intensity of the fire, and the available established on productive soils can exclude soil moisture are critical factors determin- massive sagebrush seedling establishment ing whether the plant is killed or for many years. Sagebrush seedlings gen- stimulated. erally become established on disturbed sites such as pocket gopher mounds or areas ROLE @F FIRE IN MAINTAINING trampled by wildlife or domestic livestock. MEADOW AND SAGEBRUSH- Near Blacktail Butte, on productive soils DOMINATED HABITATS where sagebrush is very dense at present, Lodgepole pine is invading sagebrush geologist Frank Bradley of the 1872 Hay- flats on alluvium along the margins of den survey party reported that “large areas forested moraines in Grand Teton National of sage had been burned off, and the grasses Park. In the Gros Ventre Valley, a similar had grown up densely, forming fine pas- invasion is occurring at forest margins. In turage and on these we again encountered the Jackson Hole area, recurrent fire seems antelope. . .” (Bradley, 1873). Compari- to have been an important factor in deter- sons of scenes in old photographs with ring substantial invasion of lodgepole pine present conditions indicate little change in into some sagebrush sites. Soil-moisture sagebrush cover on very coarse soils but a conditions in sagebrush flats preclude es- considerable increase in cover of sagebrush tablishment of lodgepole seedlings except in and other shrubs on finer-textured soils rare favorable years with abundant mois- with greater water-holding capacity. We ture. Oswald (1966) found that the years attribute the increase in shrub cover pri- 1912, 1925, and 1927 were exceptionally marily to natural plant succession in the favorable for establishment of lodgepole absence of fire. In many localities of the seedlings in the alluvial sagebrush areas west, and probably in portions of Jackson along the forest margins in Grand Teton Hole, heavy range use by cattle or sheep National Park. When a fire kills trees in has been a contributing factor in shrub in- this situation, it may take years for them 440 LOOPE AND GRUELL to reinvade. In contrast, sagebrush reinva- adapted to habitats in early stages of plant sion can be very rapid. succession. When a lodgepole pine seed source is de- One of the more obvious changes in wild- ficient, as it may be for high elevation life habitat due to fire exclusion is the loss burns, subalpine fir, Engelmann spruce, and of much willow and aspen habitat once whitebark pine become established in used by beaver. Old, barely discernible burned areas. Many such burns are above dams and gnawed stems attest to a more the elevational limits of lodgepole or are widespread occurrence of these animals in on unfavorable substrates for its establish- the past. Now that most vegetation is in ment. Reforestation of high-altitude areas a late stage of succession, the once abun- takes place very slowly. The importance of dant food source has greatly dec!ined, and fire in maintaining open areas at high ele- so has the beaver, vations in western United States is we11 On the other extreme, the evidence documented (Stahelin, 1943; Fonda and strongly suggests that have bene- Bliss, 1969; Billings, 1969). However, other fited from fire suppression. From food- factors are often involved in the mainte- habitat studies in the west, it is well estab- nance of subalpine meadows. Despain lished that big sagebrush is very often tie (1971) has emphasized the importance of staple food item in the winter diet of mule edaphic factors in maintaining meadows in deer. Big sagebrush is highly vulnerable to the . Most subalpine fire and was generally of a low density and meadows in the Jackson Hole area appear scattered occurrence on winter ranges in to be maintained by edaphic conditions, for former years. With exclusion of fire, the little conifer invasion is occurring. Massive density of sagebrush greatly increased. meadow invasions are occurring in parts of Some other preferred shrub species also in- Yellowstone National Park, however. creased in availability. The widespread in- crease in the critical carrying capacity of INFLUENCE OF FIRE ON winter range appears to be the principal WILDLIFE HABITAT reason for mule deer population increases The vegetation mosaic produced by a over pristine levels. natural fire regime results in a diverse Traditionally, moose have been asso- array of wildlife habitats. When fire is ex- ciated with seral habitats. Interestingly cluded from the landscape, plant succession enough, the moose population in northwest- leads to a greater expression of “climatic ern Wyoming was very low during the 19th climax” vegetation than would otherwise be century, when frequent fires maintained present. Research in other parts of the west much early successiona vegetation. The has suggested that early successional stages moose population in Jackson Hole has ac- favor most wildlife species. tually increased with advancing succession. The infrequency of fire in the Jackson Several contributing factors are undoubt- Hole area during the past 75 yr makes a edly involved, but a primary influence ap- local interpretation of fire’s influence on pears to be an unprecedented increase in wildlife habitat difficult. There is a critical subalpine fir-the staple food item in the need for recently burned areas for study diet of a large segment of the moose popu- purposes. Until we have new burns to lation. In coniferous forests of Jackson study, interpretations can only be made Hole, the present carrying capacity for from inference. Only a few tentative gen- moose is apparently higher than when eralizations are warranted by the available much of the forest was in early stages of evidence. However, logic suggests that succession. wildlife populations which evolved under Advanced succession has been favorable the influence of periodic fire would be best to cavity-nesting birds such FIJI the tree ECOLOGICAL ROLE OF FIRE, NORTHWESTERN WYOMING 441

swallow, mountain bluebird, house wren, large portion of the forest, including mountain and blackcapped chickadees, yel- wilderness and wildlife habitat, fires low-bellied sapsucker, woodpeckers, and would be allowed t,o burn within pre- red-shafted flicker. These species utilize old determined limits. aspen, which are very often rotten in the 2. Prescribed burning of certain areas center and susceptible to the boring of the where there is a growing need to rc- sapsucker, woodpecker, or flicker. Oppor- turn the vegetation to early succes- tunities for cavity nesting in young aspen sion. Candidate types would be livc- stands are minimal. stock and elk ranges where sagebrush has invaded grassland and aspen is de- MANAGEMENT IMPLICATIONS teriorating. If weather conditions per- mit, a 125-hectare prescribed burn of Te ton National Forest this t.ype is scheduled for the Gros Fire is a natural phenomenon and has Ventre winter range in 1973. High been the agent responsible for periodic re- priority for prescribed burning should newal of the vegetative cover. Strict control also be made for er&ensive conifrr of fire during the past three-quarters of a types where there is a need to break century has largely eliminated its natural up the fuel and thereby reduce the role in ecosystems of the Jackson Hole possibility of large-scale fires. area. Some of the environmental effects and The foregoing approach to fire managemeut management implications of this long- with attendant smoke carries with it the in- standing policy are just now surfacing. We evitable concern over air pollution. The are now beginning to understand that general public must be made aware t,hat changing the effect of one natural ferce wood smoke composed principally of car- (fire) in the ecosystem has caused the sys- bon dioxide and water has been a natural tem to respond in certain unanticipated part of the Rocky Mountain ecosystem. In ways. We have found that the energy earlier years, a pall of wood smoke hung buildup in the form of accumulated forest over the countryside quite frequent,ly dur- litter and some brush species that are ing the summer months. Being a natural favored by lack of fires (big sagebrush) is phenomenon and inevitable, it must be ac- causing the job of fire control to become cepted as a natural part of the ecosystem. more difficult and costly. We also find that the change in natural plant succession due Grand Teton National Park to lack of fire has been detrimental to some The National Park Service recognizes t,hc wildlife species, whereas others appear to need for allowing fire to play a seminatural have been favored. It appears that a close role in Grand Teton National Park if the reexamination of our fire policies is now in management goal of maintaining or restor- order. ing ecosystems to as pristine a state as pos- Our investigation suggests that the ini- sible is to be realized. During the winter tiation of a more flexible program-from of 1971-1972 a fire-vegetation management, that of strict fire control to one of fire plan was formuiated for the park and an management--is in order. Appropriate environmental impact statement was writ- actions that would help bring fire back ten to cover the effects of its implementa- into the ecosystem appear to include: tion. The plan divides the park into four 1. Delineation of the forest into fire- management zones: management zones. In some zones Zone I----All naturally caused fires al- such as valuable timber lands and lowed to burn except under exceptional cir- recreation and developed areas, strict cumst,ances (human life endangered, excep- fire control would be practiced. On a tional opportunity for spread to other 442 LOOPE AND GRUELL areas, etc.). Man-caused fires will be BOUTELLE, F. A. (1890). “Report of the Superin- extinguished. tendent of Yellowstone National Park.” U.S. Government Printing Office, 23 p. Zone II-Naturally caused fires allowed BRIDLEY, F. J. (1873). “Report of Frank H. Brad- to burn or not allowed to burn according ley, Geologist.” U.S. Geological Survey of the to judgment of a fire-management commit- Territories, U.S. Government Printing Office. tee (depending on fire-hazard conditions). BRANDEGEE, T. S. (1899). Teton forest reserve. Prescribed fires will probably be necessary House Document No. 6, 55th Congress, 34d here. session. Serial 3763. Washington, D.C. BROWN, D. (1971). “Bury My Heart at Wounded Zone III-All fires extinguished unless Knee.” Holt, Rinehart, & Winston, 487 p. they can be managed as controlled burns. BUXTON, E. N. (1893). ‘Short Stalks; or Hunting Prescribed fire will be used here to approxi- Camps.” Edward Stanford, London. mate a natural fire frequency. COLE, G. F. (1969). “The Elk of Grant Teton Zone IV-Developed areas. Manipula- and Southern Yellowstone National Parks.” Un- published manuscript. tion (logging, planting, etc.) used here to DE LACY, W. W. (1876). A trip up the South approximate effects of fire or managed for Snake River in 1863. Historical Society of Mon- aesthetically pleasing appearance. tana Contributions I, 113-118. Additional knowledge of fire behavior in DESPAIN, D. G. (1971). The vegetation of the the area will be necessary before full imple- Bighorn Mountains of Wyoming in relation to substrate and climate. Ph.D. thesis, University mentation can be attained. Whenever late- of Alberta, 136 p. summer weather conditions permit, two DOANE, G. C. (1877). “Expedition of Lieutenant prescribed burns are planned-one in a G. C. Doane-Fort Ellis, Montana, to Fort Douglas fir stand and one in an aspen-sage- Hall, . Oct. 11, 1876, to Jan. 4, 1877.” brush area. The primary purpose of these US. Department of the Interior, National Park Service. Photocopy of typed manuscript, 41 p. initial prescribed fires will be for research, FARMER, R. E. (1962). Aspen root sucker forma- for obtaining expertise in prescribed burn- tion and apical dominance. Forest Science 8, ing, and for public education. Also, starting 403-410. in the summer of 1972, lightning-caused FONDA, R. W., AND BLISS, L. C. (1969). Forest fires will be allowed to burn in the Teton vegetation of the montane and subalpine zones, Range (Zone I). In future years, prescribed Olympic Mountains, Washington. Ecological Monographs 39, 271301. burning for actual vegetation management FOWELLS, H. A. (1965). Silvics of forest trees is planned. of the United States. U.S Department of Agri- culture, Forest Service, Agricultural Handbook REFERENCES No. 271, 762 p. BAILEY, R. G. (1971). Landslide hazards related FRISON, G. C. (1971). Prehistoric occupation of to land use planning in Teton National Forest, the Grand Teton National Park. Naturarist 22, northwestern Wyoming. 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