United States 

Department of 

Agriculture 

Forest Service 

General Technical 

Report 

PNW-GTR-535 

April 2002 

Multiresource Effects of 

a Stand-Replacement 

Prescribed Fire in the Pinus 

contorta-Abies lasiocarpa 

Vegetation Zone of Central 

Washington 

Arthur R. Tiedemann and Paul M. Woodard 
Authors	 Arthur R. Tiedemann is a scientist emeritus, U.S. Department of Agriculture, Forest 

Service, Pacific Northwest Research Station, 1401 Gekeler Lane, La Grande, OR 

97850; and Paul M. Woodard is a professor, Department of Renewable Resources, 

University of Alberta, 751 General Services Bldg., Edmonton, Canada T6G 2H1. 
Abstract Tiedemann, Arthur R.; Woodard, Paul M. 2002. Multiresource effects of a 

stand-replacement prescribed fire in the Pinus contorta-Abies lasiocarpa vegetation 

zone of central Washington. Gen. Tech. Rep. PNW-GTR-535. Portland, OR: U.S. 

Department of Agriculture, Forest Service, Pacific Northwest Research Station. 26 p. 

A stand-replacement prescribed fire in an over-mature lodgepole pine (Pinus contorta 

Dougl. ex Loud.)-subalpine fir (Abies lasiocarpa (Hook.) Nutt.) stand (snag area) 

and in a mature lodgepole pine thicket (thicket area) resulted in lower plant diversity 

within the first year after burning, and as fire energy outputs increased, postburn 

plant cover and diversity decreased. There was no reestablishment of the original 

plant cover where total heat output exceeded 100 000 kcal/m2

. Apparently, most 

plants in this habitat were not fire resistant. Postfire recovery appears to depend on 

immigration of seeds from adjacent unburned areas or on seeds and rhizomes that 

survive on unburned microsites (refugia) within the burn. After fire, temperatures 

increased in the forest floor fermentative layer (FL) (10 to 19 °C) and upper 10 cm 

of the soil layer (SL) (3 to 7 °C) on several dates in summer 1976. Increased pH 

levels in FL (about 2 units) and SL (about 0.5 unit) after burning provided an 

improved environment for bacterial development, and counts of total bacteria and 

proteolytic bacteria both increased. Both nitrogen fixation and nitrification were 

increased after burning. Despite the apparent increase in microbiological activity, 

microbial respiration declined after burning—apparently because of reduced forest 

floor organic carbon energy reservoir. Diversity of birds increased the year after 

burning. New species of birds included hairy woodpecker (Picoides villosus), 

black-backed woodpecker (Picoides arcticus), three-toed woodpecker (Picoides 

tridactylus), common flicker (Colaptes auratus), and mountain bluebird (Sialia 

currucoides). Numbers of needle-foraging species, such as Townsend’s warbler 

(Dendroica townsendi ), hermit thrush (Catharus guttatus), golden-crowned kinglet 

(Regulus satrapa), and western tanager (Piranga ludoviciana), declined or were 

absent after fire. Responses of small mammals to fire were not definitive, but there 

was a marked decline in Townsend’s chipmunk (Tamias townsendii) after burning. 

In the first year after burning, forage for elk (Cervus elaphus) in the burned area 

was higher in crude protein than in unburned areas, but low productivity and 

distance from water diminished the value of the burned area for elk. 

Keywords: Forest succession, forest floor, understory vegetation, fuels, soil physical 

properties, wildlife, snags, downed wood, microbial populations, nitrification, nitrogen 

fixation, small mammals, birds, elk. 
Summary In September 1975, a prescribed fire was conducted in an over-mature lodgepole 

pine (Pinus contorta Dougl. ex Loud.)-subalpine fir (Abies lasiocarpa (Hook.) Nutt.) 

stand (snag area) and in a mature lodgepole pine thicket (thicket area). The purpose 

of the burn was to determine if a stand-replacing type of prescribed fire would 

enhance forage productivity and quality, and result in increased elk (Cervus 

elaphus) use. The area to be burned was a traditional summer elk range that was 

becoming dominated by tree cover. The burn also would provide an opportunity to 

examine the effects of fire on the indigenous plants, birds, and small mammals. 

Quantifying combustion of downed roundwood fuels and organic soil material would 

enable a better understanding of the effects of fire on various physical, chemical, and 

microbiological characteristics of the forest floor and soil. 

The prescription was designed to top-kill the existing overstory plant cover and 

eliminate surface fuels (less than 2 m tall) while containing fire within designated 

boundaries. Control areas and preburn measurements were used to assess fire 

effects. In the snag area, the fire crowned and top-killed virtually all of the overstory 

plants and reduced the surface fuel loading by 71 percent. In the thicket area, 

where crowning did not occur, overstory canopy cover was reduced by 18 percent 

and downed and dead fuels by 52 percent. 

Fire resulted in lower plant diversity within the first year after burning. As fire energy 

outputs increased, postburn plant cover and diversity decreased. Specifically, there 

was no reestablishment of the original plant cover where total heat output exceeded 

100 000 kcal/m2

. Apparently, most plants in this habitat were not fire resistant. 

Postfire recovery appears to depend on immigration of seeds from adjacent 

unburned areas or on seeds and rhizomes that survived the fire in unburned 

microsites (refugia) within the burn. There were exceptions; most notable was 

the dominance of dragonhead (Dracocephalum parviflorum Nutt.) after the fire. 

In general, plant regeneration on the site likely will be a lengthy process. 

Fire resulted in an increase in temperatures of the forest floor fermentative layer 

(FL) (10 to 19 °C) and upper 10 cm of the soil layer (SL) (3 to 7 °C) on several 

dates in summer 1976. Increased pH levels in FL (about 2 units) and SL (about 0.5 

unit) after burning provided an improved environment for bacterial development. 

Counts of total bacteria and proteolytic bacteria increased. Both nitrogen fixation and 

nitrification were increased after burning. Despite the apparent increase in microbiological activity, microbial respiration declined after burning—apparently because of 

reduced forest floor organic carbon energy reservoir. 

Diversity of birds increased the year after burning. New species of birds in postburn 

surveys included hairy woodpecker (Picoides villosus), black-backed woodpecker 

(Picoides arcticus), three-toed woodpecker (Picoides tridactylus), common flicker 

(Colaptes auratus), and mountain bluebird (Sialia currucoides). Numbers of 

needle-foraging species, such as Townsend’s warbler (Dendroica townsendi), hermit 

thrush (Catharus guttatus), golden-crowned kinglet (Regulus satrapa), and western 

tanager (Piranga ludoviciana), declined or were absent after fire. Effects of fire on 

most small mammals were not definitive, but there was a marked decline in Townsend’s 

chipmunk (Tamias townsendii ). In the first year after burning, forage for elk (Cervus 

elaphus) in the burned area was higher in crude protein than in unburned areas, but 

low productivity and distance from water diminished the value of the burned area for elk. 
Introduction In the 1970s, there was substantial interest among wildland managers in central 

Washington to devise ways to improve the carrying capacity of summer range for 

elk (Cervus elaphus) in remote forest stands in the Abies lasiocarpa zone (Franklin 

and Dyrness 1988) of the Cascade Range. Tree stands in this area are dominated 

by subalpine fir (Abies lasiocarpa (Hook.) Nutt.) and lodgepole pine (Pinus contorta 

Dougl. ex Loud.). For the most part, the commercial timber value of these stands was 

considered too low to justify costs of building roads to extract the timber. Densely 

stocked forest and large quantities of downed woody material severely restrict forage 

production and animal movement. Prescribed burning was proposed as a method 

for converting the tree-dominated stands to habitats suitable to support large herbivores. It was determined at the outset that the fire would need to be sufficiently 

severe to top-kill the existing overstory and remove a large quantity of the large 

downed logs and other debris. Essentially, this would mean a stand-replacement fire. 

Managers were concerned that this type of burn may have adverse effects on other 

resources that could outweigh benefits associated with improvement of forage and 

habitat for elk. Many of these effects were largely unknown. The U.S. Department of 

Agriculture, Forest Service (FS), Pacific Northwest (PNW) Research Station viewed 

this as an opportunity to gain a better understanding of the effects of a controlled, 

stand-replacement fire on a variety of resources on the east side of the Cascade 

Range. In a partnership with the Ellensburg Ranger District of the Wenatchee 

National Forest, Pacific Northwest Region of the FS, a plan was developed to conduct 

a prescribed burn in a 10.9-ha area with two different-aged contiguous stands. The 

younger stand, referred to as the “thicket area,” was dominated by mature lodgepole 

pine with subalpine fir and Engelmann spruce (Picea engelmannii Parry ex Engelm.) 

in the intermediate and suppressed crown classes (Woodard 1977, 1993b). The 

older stand, referred to as the “snag area,” was over-mature lodgepole pine with 

large, live subalpine fir and Engelmann spruce as codominants in the overstory 

canopy. The cooperation of the staff at the University of Washington, Seattle; 

Washington State University, Pullman; and Central Washington University, 

Ellensburg, was obtained in an attempt to capitalize on the amount and quality of 

information to be realized from what eventually would be a moderately expensive 

study. The PNW Research Station at Wenatchee provided funding to each university 

through cooperative aid contracts. Results of one study were published by scientists 

at the PNW Research Station (Fowler and Helvey 1978). In addition, three M.S. theses 

and one Ph.D. dissertation were completed based on research conducted before 

and after the prescribed fire. Information from the Ph.D. dissertation of Woodard 

(1977) was the basis for three additional publications (Woodard 1983, 1993a, 1993b). 

The five studies were designed to capture information pertaining to (1) the thermal 

regime and total heat output at the time of burning; (2) the assessment of physical 

effects of the fire on standing trees and the downed fuels; (3) the first-year response 

of plants to a stand-replacing crown fire; (4) the changes in forest floor, to include 

soil properties and microbial populations and processes; (5) the effects of burning 

on birds and small mammals; and (6) the changes in forage production, forage quality, 

and elk use. Although this research plan did not include all factors naturally affected 

by a fire, it served as a start to understanding the natural role of fire in this subalpine 

forest zone and to test a fire prescription that may serve managers in the future. 

1

Methods of Study 

The Study Area 

It seems timely to summarize information on the effects of this prescribed fire 

because of the emphasis on restoring fire to forests of the interior Pacific Northwest 

(Everett and others 1993, Hardy and Arno 1995, Wickman 1992) and concerns 

about its effects on all resources (Tiedemann and others 2000). Therefore, the goal 

of this paper is to synthesize the information in these theses and publications on the 

effects of prescribed fire in the lodgepole pine-subalpine fir “snag” and “thicket” burn 

areas. Although a stand-replacement fire is not the goal for most prescribed burns, it 

must be anticipated that, despite careful planning, this type of fire is likely to occur in 

some prescribed burns. This compilation is meant to provide much needed information 

for planning effects of stand-replacement prescribed burning on forest resources. 

This type of information is important for developing strategies for managing forest 

resources with prescribed fire (Tiedemann and others 2000). Another purpose of 

publishing this paper is to make people aware of the wealth of information available 

from the original Ph.D. dissertation (Woodard 1977) and three M.S. theses (Boltz 

1979, Hanson 1978, Husby 1978). 

An area of the eastern slope of the Cascade Range called Table Mountain, located 

about halfway between Wenatchee and Ellensburg, Washington (USA), was the site 

chosen for the 10.9-ha prescribed burn. It is about 2 km from Haney Meadow up a 

steep mountain game trail in S. 24, T. 21N, R. 18E, Willamette P.M. (latitude 14° 18’ N., 

longitude 120° 32’ W). The site was representative of many lodgepole pine-subalpine 

fir stands in the Table Mountain area. The disadvantage of inaccessibility for collection 

of data was offset by the advantage of lack of disturbance by recreationists using 

nearby Haney Meadows. Remoteness also provided managers with an indication of 

the difficulty and costs of conducting prescribed burns in remote sites, but those 

issues are not addressed in this paper. 

Woodard (1977, 1993a, 1993b) provided a comprehensive description of the vegetal 

and physical characteristics of the study area. Elevation varied narrowly between 1706 

and 1761 m on a gentle southwest slope. Soils are described in detail by Woodard 

(1977) from descriptions provided by McColley (1976). Soils are not named or 

classified. Surface soils are derived from volcanic ash over residuum of basalt 

bedrock. Soils vary from 10 to 50 cm in the thicket area and from 40 to 100 cm in the 

snag area. 

According to Woodard (1977), climate is typical of the subalpine zone with mean 

minimum January temperature about -8 °C and mean maximum July temperature of 

28 °C. Most (70 percent) of the annual 80 to 160 cm of precipitation occurs as snow 

between October and March (Fowler and Helvey 1978). 

For the thicket area, the live tree basal area and density were 72.1 m2

/ha and 1,505 

trees per hectare, respectively. Live tree basal area and density in the snag area 

were 55.8 m2/ha, and 1,368 trees per hectare, respectively. Woodard (1977) 

described the thicket area as a well-differentiated stand. The canopy of the closed 

stand was dominated by lodgepole pine, with subalpine fir and Engelmann spruce 

as codominants in the intermediate crown class. The stand in 1975 was predominately 

105 years old, although there were trees of various ages throughout the area. 

Subalpine fir was the only tree species regenerating in the understory. Elk sedge 

(Carex geyeri Boott), arnica (Arnica cordifolia Hook. and A. latifolia Bong.), Hood’s 

sedge (Carex hoodii Boott), lupine (Lupinus polyphyllus Lindl.), and low whortleberry 

2

(Vaccinium myrtillus L.) were the predominant understory plant species. In the snag 

area, standing snags of lodgepole pine were obvious in the overstory canopy 

suggesting that it was an over-mature lodgepole pine stand (Woodard 1977). 

The oldest trees established 305 years before 1975. Subalpine fir and Engelmann 

spruce were codominants in the overstory canopy and appeared in the intermediate 

strata of the canopy as well. Again, only subalpine fir was commonly found in the 

understory layer. The principal understory plants in this stand were elk sedge, 

arnica (A. latifolia), and mosses (Polytrichum commune Hedw. and Racomitrium 

canescens var. ericoides (Brid.) B.S.G.). 

The fire history of both stands was determined by Woodard (1977, 1993a). He used 

fire scars, increment cores, and bole disks from 195 trees to uncover the historical 

role of fire in the area. He documented the occurrence of three small fires–1835, 

1850, and 1890–in this 10.9-ha treatment area (Woodard 1993a). He concluded that 

“what appeared to be two even-aged lodgepole pine stands with subalpine fir and 

Engelmann spruce in intermediate height classes, were complex, multiaged, mixedspecies stands where all species were common in all age classes and all species 

were capable of surviving and reestablishing after fire” (Woodard 1993a). 

The study area provided habitat for various birds including mountain chickadee 

(Parus gambeli ), red-breasted nuthatch (Sitta canadensis), golden-crowned kinglet 

(Regulus satrapa), ruby-crowned kinglet (R. calendula), yellow-rumped warbler 

(Dendroica coronata), Townsend’s warbler (D. townsendi ), varied thrush (Ixoreus 

naevius), hermit thrush (Catharus guttatus), red crossbill (Loxia curvirostra), 

Cassin’s finch (Carpodacus cassinii ), evening grosbeak (Coccothraustes 

vespertina), Stellar’s jay (Cyanocitta stelleri ), chipping sparrow (Spizella passerina), 

and various hawks and owls (Hanson 1978). Elk was the primary large herbivore in 

the study area. Small mammals included yellow pine chipmunk (Tamias amoenus), 

Townsend’s chipmunk, red-backed vole (Clethrionomys gapperi), deer mouse 

(Peromyscus maniculatus), northern flying squirrel (Glaucomys sabrinus), goldenmantled ground squirrel (Spermophilus saturatus), short-tailed weasel (Mustella 

erminia), shrew-mole (Neurotrichus gibbsii ), vagrant shrew (Sorex vagrans), snowshoe hare (Lepus americanus), and Douglas squirrel (Tamiasciurus douglasii ) 

(Hanson 1978). 

Cattle, sheep, and horses have used the area extensively since the time of settlement (Woodard 1977). Sheep used the area from 1907 until the FS began to 

restrict use in 1923. Between 1924 and 1972, the treatment area was part of a cattle 

allotment. After 1972, cattle use was excluded from the allotment. During the 

recorded period of use, animal use month (AUM) estimates for the entire allotment 

varied from 8,300 AUMs in 1924 to 925 AUMs in 1958 (Woodard 1977; based on 

records from the Wenatchee National Forest). In 1975, use in excess of 4,007 

AUMs was considered “overuse.” This level of use likely had an impact on fuel, 

plant cover, and soil compaction. According to FS records, elk were introduced in 

1915 (Woodard 1977). Currently, these high-elevation areas are used for calving 

and as summer and fall range for an undetermined number of elk. The length of 

time the area is used annually is determined by snow depth and hunting pressure. 

3

Table 1—Summary of fire prescriptions (proposed and actual) for the snag 

and thicket areas of the Table Mountain prescribed burn project in central 

Washingtona 

Proposed 

Parameter USFSb U of Wc Actual 

Ambient air 

temperature 

Relative humidity 

Wind (speed and 

direction) 

Days since rain 

Fuel moisture content 

Low 16 °C 

35-45% 

<8 km/hr (S) 

– 

– 

(fine) 

Method of firing 

Proposed date for 

ignition 

16-27 °C 

<40% 

5-13 km/hr (S) 

4+ days, once 

fuels cured 

10-20% 

Backfire north side with 

3-meter strips, then strip head fire 

remainder at 15-meter intervals 

16-17 °C 

19-21% 

Calm, gusts to 

26 km/hr 

(S-SW erratic) 

15 days 

13.1% 

As proposed 

Sept. 30, 1975 

1400-1600 hrs 

by using 2 crews


Mid-Sept. 1975 Mid-Sept. 1975 


Fire Prescription and 

Conduct of the Burn 

a 

b From Woodard (1977). 

USDA Forest Service. c 

University of Washington. 

The objective of the burn was to achieve a manageable crown fire, which would 

top-kill the overstory tree cover and eliminate as much of the accumulated surface 

fuels as possible without exceeding the treatment boundaries. Woodard (1977) 

detailed the fire prescription and provided a detailed description of the fuel moisture 

content and weather conditions necessary to accomplish these objectives and the 

actual conditions at the time of ignition (table 1). 

Ignition of the fire occurred on September 30, 1975, beginning at 1406 hours. 

Woodard (1977) described the procedure for ignition and progress of the fire. 

Two four-man firing crews backfired the northeast quadrant of 

the treatment area. Fire strips, 3 meters apart for the first 25 

meters, were backed into the hand-constructed firebreak, thereby 

increasing the depth of the control line. The wind was blowing 

from south to north. A 50/50 diesel oil and gasoline mixture 

4

Results 

Documented 

Characteristics 

of the Fire 

dripped from a hand-held torch was used to ignite 

down and dead fuel. After the safety zone had been 

widened, the burners proceeded to strip headfire the 

remaining area. Four men, 6-12 meters apart, dripped 

parallel lines of fire that traversed the treatment area 

from northwest to southeast. Within approximately 10 

minutes, trees were beginning to crown-out. Flame 

heights were estimated to be 38 meters by one observer 

and 15 meters over the tops of 27 meters crowns by two 

other people. 

About 1 hour and 20 minutes were required to ignite the 10.9-ha area. Although 

burnout time varied among specific locations, most of the area was in the glowing 

phase of combustion by 2300 hours on October 1, 1975. The fire smoldered 

throughout most of October 2, 1975, but risk of the fire escaping outside the 

treatment area was small, and fire crews began to pack up their equipment 

(Woodard 1977). Within less than 2 weeks of ignition, the winter snows began to 

fall and the fire was declared out. 

This is not a replicated study. Some of the statistical comparisons are pseudoreplication using sample dates as replicates to compare the control with burned areas 

and for comparisons among years. This is especially true for the soils and forest 

floor chemical and microbiological results. 

Prescribed fire in the snag area resulted in a crown fire that, with exception of one 

small spot fire, was managed within prescribed limits (Woodard 1977). Crowning 

was most prevalent in the snag area but rare in the lodgepole pine thicket area. This 

was likely a result of the differences in surface fuel loadings between the two areas. 

Fire temperatures measured by Fowler and Helvey (1978) exceeded 900 °C in the 

snag and thicket areas at the soil surface and up to 2 m height. Only one measurement point was documented for the thicket area, so the information applies mainly 

to the snag area. Temperatures of 700 °C were achieved at 3 m and in excess of 

600 °C at 5 m heights. 

Woodard (1977) found the total heat output calculated from fuel reduction measurements provided the best estimate of the thermal environment of the site. Following 

fuel sampling procedures described by Brown (1974), preburn and postburn fuel 

loads were determined for 30 permanent, random sample plots. The difference 

between these two loadings is commonly termed the “available” (Byram 1958) fuel 

loading (loss). The available fuel weight in metric tons per hectare then was 

multiplied by 450 to estimate potential heat outputs in kilocalories per square meter. 

The reader is referred to Woodard (1977) for details of this analysis and the other 

estimators used in that work. Heat outputs in the snag area ranged from 1 530 to 

1 542 420 kcal/m2

. In the thicket area, burned plots were estimated to have released 

630 to 908 865 kcal/m2

. Unburned plots were estimated to have a heat release 

value of 0 kcal/m2. Woodard (1977) suggested these levels of heat output are 

inordinately broad, but realistic for natural conditions. 

5

Effects of Fire on 

Tree Basal Area and 

Canopy Cover 

Effects of Fire on 

Accumulated Surface 

Fuels 

Vegetation Responses 

The basal area of the preburn live tree stems and the overstory canopy cover was 

reduced because of burning. In the thicket area where crowning did not occur, the 

average change in basal area of live stems was -1.5 percent, whereas the average 

percentage change in the overstory canopy cover was -18 percent. In the snag area 

where crowning had occurred, the basal area was reduced an average of 15 percent, 

and the average change in the overstory canopy cover was -63 percent. 

Total accumulations of the surface fuels, including the organic matter in the forest 

floor and downed-dead tree boles, averaged 174 t/ha for the thicket area and 268 

t/ha for the snag area (Woodard 1977). Fuels were divided into two general categories: duff and downed roundwood. The downed roundwood weight was determined for wood diameters smaller than 0.63 cm, 0.63 to 2.5 cm, 2.5 to 7.6 cm, and 

larger than 7.6 cm. Fuels between 0 and 7.6 cm in diameter are commonly called 

fine fuels and contribute significantly to fire spread, whereas fuels larger than 7.6 cm 

diameter are coarse fuels and are important in burn-out or residence time. Duff is 

the common name used to describe the fermentation (F) and humus (H) layers of 

the mineral soil (Hoover and Lunt 1952, Kittredge 1948). It is also known as the Oe 

and Oa horizons (Page-Dumroese and others 1991). In the thicket area, 54 percent 

of fuels were in the roundwood categories, while the remaining was duff. Roundwood 

fuels composed a much higher percentage (75 percent) of total fuels in the snag 

area. Roundwood larger than 7.6 cm in diameter accounted for most of the difference in the roundwood weight in the snag area. Woodard (1977), however, found 

that fine fuel loadings were also higher in the snag area (19.8 t/ha) when compared 

to those in the thicket area (15.7 t/ha). As expected, fuels were highly variable 

among sample points even within the same area. For example, in the thicket area, 

total fuel weight varied almost fivefold from 58 to 270 t/ha. Variation in total fuel 

weight in the snag area was even greater (sixteenfold), from 49 to 781 t/ha. 

Burning resulted in total fuel weight reductions in the thicket area that ranged from 

zero to 84 percent and averaged 52 percent (Woodard 1977). In the snag area, the 

total fuel weight reduction ranged from 13 to 97 percent and averaged 71 percent. 

As expected, the largest reductions occurred in fuel smaller than 7.6 cm in diameter—96 percent in the snag area and 70 percent in the thicket area. Reductions in 

the weight of duff loading were indicative of the extreme severity of the fire in the 

snag area and the lower severity in the thicket area. The duff weight in the snag 

area was reduced from 60 to 4 t/ha. In the thicket area, duff weight was reduced 

from 74 to 49 t/ha. More specific information about fuel reduction by individual fuel 

size class is in Woodard (1977). 

Woodard (1977, 1983, 1993b) studied plant community composition responses to 

the fire in the snag and thicket areas to determine if the prefire plant community 

composition could be used to accurately predict the composition of the postfire 

community during the first year after the burn. He used permanently located belt 

transects, established before burning, to measure onsite vegetation responses to 

fire (fig. 1). Understory vegetative cover was sparse before burning—12 percent and 

18 percent for snag and thicket areas, respectively (Woodard 1977). Highest 

understory plant diversity or species richness before burning was found in the snag 

area where 32 species were identified compared to 27 in the thicket area. However, 

heat outputs from the fire were so great in the snag area that no plants reestablished 

the first year on 90 percent of the permanently located plots (fig. 1). Actual percentage 

6

of ground covered by vegetation was reduced by burning in both snag and thicket 

areas, but of the total area available for plant occupation, a greater percentage was 

covered by postfire plant species. The number of plant species in the snag area 

was reduced from 32 to 18. This change included 7 new species, which were found 

for the first time after burning, and the absence of 21 of the original species found 

before burning. In the thicket area, the total number of plant species declined from 

27 to 24 as a consequence of burning. Five new species immigrated to the area, 

and eight prefire species were absent after burning. 

Woodard (1977) assessed the change in cover for subalpine fir, both arnica species, 

two moss species, lupine, and sedges (Carex species summed) as a function of 

heat output. Although Woodard (1977) was not able to establish statistical relations, 

the plotted data indicated that as total heat output increased, the percentage of 

cover of all seven species and species groups declined. More importantly, when heat 

output exceeded 100 000 kcal/m2

, none of the seven species or groups found on 

plots before burning had regenerated 1 year after burning. Subalpine fir was the 

species most sensitive to increasing heat output. It did not regenerate on plots that 

received more than 60 000 kcal/m2. Lupine, a rhizomatous species that was anticipated to be capable of regenerating after intense fire, had not appeared on the plots 

after 1 year when the heat output exceeded 100 000 kcal/m2. Woodard (1977) did 

find arnica (A. latifolia) and sedges on plots where heat output exceeded 100 000 

kcal/m2, but these were plots where the species were not observed before burning. 

Woodard (1977) speculated that establishment was the result of seed migration or 

germination of seed that survived burning. 

Plant species that increased most after burning were dragonhead, arnica, and moss 

(Polytrichum commune Hedw.). The response of dragonhead was of particular 

interest because it contributed most of the postburn cover in the snag area and was 

not identified in preburn surveys. It appears that this species is a strong invader of 

burned sites and is an important early seral species in these habitats. The two plant 

species that seemed most sensitive to burning were fireweed (Epilobium angustifolium L.) 

and columbine (Aquilegia flavescens S. Wats.). They were not found in either the 

snag or thicket burn areas the first year after burning. The response of other individual 

plant species to the effect of the fire are presented by Woodard (1977, 1993b). In 

general, field measurements indicated that most of the plants in this habitat were 

not fire-resistant or did not have adaptive mechanisms that enabled them to survive 

a high-intensity fire such as the one applied in this study. 

Woodard (1977, 1993b) also explored mechanisms of plant regeneration by a 

greenhouse study of plant emergence from soil cores extracted 1 week before and 

2 weeks after burning. For both snag and thicket areas, he identified 16 plant 

species on preburn soil cores before extraction. After 10 months in the greenhouse, 

21 plant species had emerged (Woodard 1993b). Seven species from preburn soil 

cores were not encountered in field surveys and seven species found in field surveys 

before core extraction were not found in either preburn or postburn soil cores during 

the greenhouse test. These results suggest that cores contained seeds of plants 

that were not found in the area before the fire was ignited. One can assume that the 

field conditions before burning were not suitable for these species or that these 

seeds may have been deposited during the previous season from plants outside of 

the treatment area. 

7 
b 

a 

Figure 1—a. Preburn 1975 vegetation transect in the snag area. Note downeddead and forest floor accumulations and sparse understory cover. b. Postburn 

1976 vegetation transect from the same photopoint portrays the aftermath of 

the fire. c-e. 1977, 1978, and 1980 photos of vegetation transect portrays the 

slow recovery of vegetation. 

8

c 

d 

e 

9

Microbiological 

Responses of the 

Fermentative Layer 

and Mineral Soil 

Neither Engelmann spruce nor lodgepole pine were found on field plots after burning 

(Woodard 1977, 1983). To test the hypothesis that seedbed suitability was the factor 

that limited recruitment of these tree species after fire, he germinated seeds of 

Engelmann spruce and lodgepole pine on (1) white filter paper watered with distilled 

water; (2) white filter paper watered with dilute (500 mL ash and 1000 mL distilled 

water) and concentrated (500 mL ash and 500 mL distilled water) ash solutions; (3) 

35 mL of washed ash solution; and (4) 35 mL untreated ash solution. Both species 

germinated in all treatments (73 to 92 percent germination). Engelmann spruce germination was not affected by any treatment. Lodgepole pine germination was reduced 

by both ash solutions and by the untreated ash solution. Even though germination 

percentages of both species were relatively high for all treatments, Englemann 

spruce has an advantage over lodgepole pine because it has higher germination 

levels and was not affected by ash solution or ash bed treatments. 

Effects of fire on pH, temperature, moisture content, total bacteria, fungi, proteolytic 

bacteria, actinomycetes, nitrification rates, nitrogen fixation, and microbial respiration in the fermentative layer (FL) of the forest floor and upper 10 cm of soil were 

determined by Husby (1978). Husby investigated effects in both thicket and snag 

areas as separate study units. The FL was defined as that organic material between 

the litter layer and mineral soil. Mineral soil layer (SL) was sampled to 10 cm depth. 

The FL, thus, included F and H (Hoover and Lunt 1952, Kittredge 1948) or Oe and 

Oa layers, respectively (Page-Dumroese and others 1991). Before the fire in 1975, 

samples were collected from four random locations in 8- by 8-m permanent plots in 

snag and thicket areas to be burned and from one control plot for each. Control 

plots were about 300 m from burned plots. Preburn samples were collected biweekly 

starting when snow was gone from plots and continuing until the site was burned. 

Postburn sampling in 1976 began as soon as snow was gone and continued 

biweekly until the first snow of the fall. In 1977, sampling started with snow elimination 

and continued at biweekly intervals until mid-July. Details of sample handling, 

preparation, and analytical procedures are in Husby (1978). He measured effects of 

burning on temperature, pH, and moisture content of the FL and SL because those 

factors govern the rate of biochemical processes. Results were tested at a probability 

level of P < 0.05 for the F-test and for comparisons of means among years at 

individual sites. Husby (1978) did not use a statistical analysis that would enable 

comparison of the control area with the burned area, assess differences among 

individual sample dates, or determine interaction of burning with sample dates or 

years. It also should be noted that statistical comparisons between 1977 and the 

previous 2 years may be compromised because sampling ended in mid-July 1977. 

Temperatures of the FL and SL did not differ significantly among years for the control 

plot (tables 2 and 3). Burning did not significantly influence the FL or SL mean 

temperatures in the thicket or snag areas. Comparison of temperatures of control 

and burn areas that Husby (1978) observed on specific dates provided a more 

comprehensive indication of temperature effects (although not statistical) than did 

comparison of mean temperatures. For the thicket area, postburn temperatures of 

the FL were relatively uniform between the control area and burn area for all sample 

dates (Husby 1978). For the snag burn area, however, temperatures of the FL were 

10 to 19 °C greater (no statistical test) than the control in mid-June and July 30 to 

August 4, 1976. Fowler and Helvey (1978) measured a 26 °C increase in maximum 

temperature (at 1 cm above the surface) after burning in 1976. In 1977, Husby 

10

(1978) found that increases for individual dates had moderated to 5 to 7 °C. 

Postburn temperatures in 1976 on individual sample dates in the SL increased by 3 

to 7 °C. Husby (1978) attributes these higher temperatures to virtually complete 

removal of all foliage and most branches from trees, removal of a large proportion of 

the litter, and the blackened remnant of the forest floor (Woodard 1977). The fire in 

the thicket area, in contrast, was not nearly as severe as in the snag area and did 

not crown out; much of the tree foliage, therefore, was left intact. Also, there was not 

nearly as complete consumption of litter and duff, and the forest floor was not as 

thoroughly blackened as in the snag area. 

Moisture contents of the FL and SL did not differ significantly among years for the 

control plots (tables 2 and 3). In the thicket area, burning resulted in increased 

moisture in the FL in 1977 and in the SL in both 1976 and 1977. Moisture content 

more than doubled between 1975 and 1977 in both layers. In the snag burn area, 

there were no significant moisture-content differences among years for the FL or SL. 

Mean pH in the snag area control did not change significantly among years in the 

FL or SL (table 2). However, in the thicket area control, there was a significant 

increase of 0.5 pH unit in both the FL and SL between 1975 and 1977 (table 3). 

Husby (1978) attributed this to natural year-to-year variation. In the thicket area, 

burning resulted in a significant increase of nearly 2 pH units in the FL and about 

0.5 pH unit in the SL. The increase was significant in both layers in 1976. In the 

snag area, burning resulted in increased mean pH in the FL for both postburn years 

(table 2). Increases in 1976 and 1977 were 2.2 and 1.8 pH units, respectively. After 

burning, the SL pH in the snag area increased significantly only in 1976, but only by 

0.4 pH unit. 

Bacteria are involved in mineralization processes, whereby nutrients such as nitrogen 

(N) and sulfur (S) are converted from an intrabiotic organic state to a mineral state, 

which is required for plant uptake. Fire has the potential to influence many factors 

that govern bacterial populations, specifically pH, moisture, temperature, and 

energy substrates. Total bacteria counts (number per gram) were not significantly 

different among years in the FL or SL of control plots (tables 2 and 3) (Husby 1978). 

Counts increased significantly from 1.9 x 106 before burning in 1975 to 1 x 108 in 

1977 after burning in the FL of the thicket area, but there was no change in counts 

of the SL (table 3). After burning, bacteria counts increased significantly in both the 

FL and SL of the snag area (table 2). In the FL, counts increased from a preburn 

level of 7.6 x 106 to 1.1 x 108 between 1975 and 1977. A significant increase in 

counts in the SL also occurred between 1975 and 1977 (1.7 x 106 to 1.8 x 107) in 

the snag area. 

Fungi were measured as part of this study because they compose the largest 

proportion of total microbial protoplasm in well-aerated soils (Alexander 1961, 

Husby 1978). They also are known to be important in the decomposition of 

substances, such as lignin, which are resistant to bacterial degradation (Alexander 

1961, Husby 1978). Generally, fungi are more common in acidic soils, whereas 

bacteria are more abundant in soils that are nearly neutral for pH. Husby (1978) 

found that fungi displayed a large natural variation among years as indicated by 

significant differences among years on the control plots (tables 2 and 3). It appeared 

that fire influenced only fungi in the SL of the thicket burn area (table 3). However, 

11 
Table 2—Comparisons among years of several physical and microbiological characteristics in the 

fermentative layer (FL) and upper 10 centimeters of soil (SL) for snag burn (SB) and control areas (C) 

of the Table Mountain prescribed burn project in central Washington1 

Characteristic 

FL SL 

1975 1976 1977 1975 1976 1977 

pH 

Temperature 

(°C) 

Moisture content 

(percent) 

Total bacteria 

(number/gram) 

Fungi 

(number/gram) 

Proteolytic bacteria 

(number/gram) 

Actinomycetes 

(number/gram) 

N2 fixation 

(nanomoles 

ethylene/day) 

Microbial respiration 

(mg CO2/day) 

C 5.2a2 5.1a 5.2a 5.5a 5.4a 5.5a 

SB 4.9a 7.1a 6.7b 5.4a 5.8b 5.9b 

C 8.4a 6.9a 5.7a 7.1a 5.7a 4.7a 

SB 9.2a 13.9a 10.9a 7.2a 9.7a 8.9a 

C 35.5a 61.5a 55.6a 18.8a 20.8a 19.6a 

SB 31.0a 14.0a 25.0a 19.7a 23.6a 23.0a 

C 6.8x106

a 1.6x107a 1.6x107a 4.3x106a 3.8x106a 3.8x106a 

SB 7.6x106a 5.9x107b 1.1x108c 1.7x106a 9.7x106ab 1.8x107bc 

C 4.8x105

a 1.6x106b 6.5x105a 1.9x105a 5.1x105a 2.5x105a 

SB 3.5x105a 3.3x105a 4.9x105a 1.8x105a 4.8x105a 7.1x105a 

C 1.2x106

a 3.1x106a 4.8x107a 5.4x105a 7.8x105a 4.5x107a 

SB 5.7x105a 1.8x106a 1.5x107b 3.2x105a 7.0x106a 2.3x106b 

C 2.4x105

a 1.0x106a 5.8x105a 1.2x105a 1.3x106a 1.4x105a 

SB 1.2x105a 4.2x105a 1.0x104a 5.2x104a 1.9x105b 1.3x106c 

C 1.6a .8a 1.4a <.1a 0a <.1a 

SB .5a 14.5a 32.2b <.1a .1a .9 

C 3.0a 3.5a 3.2a .9a .9a .7a 

SB 2.7b 1.2a 1.4a .9a .8a .7a 

1 From Husby (1978).
 2 Values in a row within an individual FL or SL category with the same lowercase letter are not significantly different among years at P < 0.05.


because of the natural variation that was of similar magnitude among years at the 

control area, the effect of fire cannot be statistically determined. 

Proteolytic bacteria are important in decomposition of proteins to amino acids 

(Alexander 1961). Husby (1978) observed a significant year-to-year variation in the 

control plot FL of the snag area (table 2). Proteolytic bacteria in 1977 were more than 

10 times greater than in 1975 or 1976. There were significant increases in proteolytic 

bacteria as a result of burning in the SL of the thicket area (more than tenfold increase) 

and in the FL of the snag area (slightly less than tenfold increase) (tables 2 and 3). 

Husby (1978) found that actinomycetes did not vary significantly among years on 

12

Table 3—Comparisons among years of several physical and microbiological characteristics in the 

fermentative layer (FL) and upper 10 centimeters of soil (SL) for thicket burn (TB) and control areas 

(C) of the Table Mountain prescribed burn project in central Washington1 

Characteristic 

FL SL 

1975 1976 1977 1975 1976 1977 

pH 

Temperature 

(°C) 

Moisture content 

(percent) 

Total bacteria 

(number/gram) 

Fungi 

(number/gram) 

Proteolytic bacteria 

(number/gram) 

Actinomycetes 

(number/gram) 

N2 fixation 

(nanomoles 

ethylene/day) 

Microbial respiration 

(mg CO2/day) 

C 4.4a2 4.4a 4.9b 5.1a 5.3a 5.8b 

TB 4.5a 6.4b 6.3b 5.2a 5.9b 5.9b 

C 8.9a 6.8a 6.4a 7.8a 5.9a 4.9a 

TB 9.2a 6.8a 5.8a 7.6a 5.9a 3.9a 

C 59.3a 100.1a 62.8a 25.7a 30.6a 29.4a 

TB 55.0a 68.0a 116.2b 23.7a 37.4b 63.0c 

C 1.6x107

a 1.7x107a 6.2x106a 4.4x106a 4.8x106a 9.7x106a 

TB 9.9x106a 8.7x107ab 1.0x108b 3.0x106a 1.8x107a 8.2x106a 

C 1.4x106

a 3.7x106a 3.7x106a 2.4x105a 2.5x105a 7.0x105b 

TB 1.8x106a 8.6x105a 1.3x106a 2.6x105a 1.6x105a 5.3x105b 

C 2.8x106

a 6.5x106a 2.5x107a 6.9x105a 8.6x105a 3.3x106a 

TB 3.1x106a 1.4x107a 3.4x107a 3.2x105a 2.4x106a 5.4x106b 

C 1.3x105

a 1.1x106a 5.8x105a 1.3x105a 2.5x105a 2.5x105a 

TB 7.2x104b 4.3x105a 3.7x105a 5.9x104a 1.8x105a 3.3x105a 

C 1.4a .6a 43.3a <.1a .2a 12.6b 

TB 1.2a 15.1a 58.8b .1a 2.0a 39.1 

C 3.3a 4.3a 3.0a 1.0a 1.0a .7a 

TB 3.4b 1.8a 1.8a .8a .9a .7a 

1 From Husby (1978).
 2 Values in a row within an individual FL or SL category with the same lowercase letter are not significantly different among years at P < 0.05.


the control plots (tables 2 and 3). There were significant increases in actinomycetes 

in both the FL and SL of the thicket area in response to burning (table 3). In the 

snag area, actinomycetes increased significantly after burning only in the SL (table 2). 

Effects of fire on N fixation were measured to determine whether N replenishment of 

the soil/forest floor system was enhanced after fire. These high-elevation sites typically 

are deficient in N (Tiedemann and Klock 1977), and losses from fire can be 

substantial (Raison and others 1984). Inputs from precipitation are small, commonly 

less than 2 kg/ha per year (Tiedemann and others 1978). Thus, fixation is probably 

the primary means of N replenishment (Binkley 1986). Nitrogen fixation as 

13

Effects of Fire on Bird 

and Small Mammal 

Populations 

measured by acetylene reduction was less than 2 and less than 0.1 nanomoles 

of ethylene per day in the FL and SL, respectively, of the snag area control plot (for all 

years). Levels were similar before burning on thicket area and snag area burn plots 

(tables 2 and 3). However, N fixation in the control plot of the thicket area increased 

substantially in both the FL and SL during the 1977 sampling season (table 3). 

Nitrogen fixation increased significantly (14 to 28 times) after burning in the thicket 

area and snag area in the FL and SL. The increase was significant only for 

comparisons of 1975 and 1977. 

Nitrification is part of the N mineralization process whereby plant proteins are 

converted from ammonium-N to nitrate-N. These are the two forms of N taken up by 

plants and used in formation of proteins and metabolites. Husby (1978) measured 

nitrate production (micrograms per gram of FL or SL after 15-day perfusion) before 

and after burning (only in the snag area) as an indication of the effect of fire on rate 

of nitrification. Nitrate accumulation before burning in the FL and SL was less than 

0.01 μg/g. For the mid-June and early July samples in 1976, there was no apparent 

effect of fire on nitrification. Nitrification, however, rose sharply in both layers after 

July 1, 1976. Nitrate levels observed July 15 through October 20, 1976, in the FL 

ranged from 10.5 to 61 μg/g. In the SL, levels for the same period ranged from 1.7 

to 16.7 μg/g. Nitrification was still elevated (7.5 to 23 μg/g) in the FL during the 1977 

sample period. In the SL, nitrate production in 1977 was sharply lower than 1976 

(0.1 to 0.2 μg of nitrate per gram of soil). 

Measurements of microbial respiration (mg of carbon dioxide per day) are indicative 

of microbial processes and adequacy of relatively available energy sources for 

microbial populations (Hu and others 1972). In Husby’s (1978) study, respiration 

was used to provide an indication of the net effects of burning on microbial populations and processes. Microbial respiration in the FL was about three times greater 

than in the SL (tables 2 and 3). In the thicket area control, there was significant 

yearly variation (table 3). Mean microbial respiration declined from 4.3 to 3.0 mg 

of carbon dioxide per day between 1976 and 1977. Burning caused a significant 

reduction in mean microbial respiration in the FL at the snag and thicket areas 

(tables 2 and 3). In the thicket area, mean FL respiration declined from 3.4 to 1.8 mg 

of carbon dioxide per day between 1975 and 1977 (table 3). In the FL of the snag 

area, the decline was from 2.7 to 1.4 mg of carbon dioxide per day (table 2). 

Hanson (1978) conducted preburn and postburn censuses of bird and small mammal 

populations. The snag burn area and thicket burn area were combined into a single 

unit (snag-thicket). Hanson’s (1978) unburned control was near Haney Meadow 

about 1.6 km north-northwest of the snag-thicket burn area. Forest composition and 

structure of the control were similar to those of the snag-thicket burn area. 

Birds—Hanson (1978) made same-day measurements in the snag-thicket burn 

area and in the control area. Both areas were gridded by Cartesian coordinates to 

provide permanent reference points. Bird density by species was determined on 

each area between 0530 and 1100 hours on seven dates June through July 1975, 

and May through July 1976, except that in 1976, there were only six census dates in 

the control area (Hanson 1978). A total of 244 and 252 points were sampled in the 

snag-thicket burn area in 1975 and 1976, respectively. In the control area, 151 and 

111 points were sampled in 1975 and 1976, respectively. 

14

Territories of male birds were determined by plotting the locations of singing male 

birds over space and time onto composite maps (Hanson 1978). Diversity indices 

were calculated from counts of all birds recorded using the area. Relative abundance 

summed over time was used to give an index of community bird diversity. Refer to 

Hanson (1978) for detailed descriptions of methods used. Species for which densities were not determined as territories, but for which relative abundances were used 

in determination of diversity indices included varied thrush, red crossbill, Cassin’s 

finch, evening grosbeak, Stellar’s jay, gray jay (Perisoreus canadensis), chipping 

sparrow, and hawks and owls. 

During 1975, Hanson (1978) found seven species of breeding birds in the control 

area and eight species in the area to be burned (tables 4 and 5). Townsend’s warblers, 

golden-crowned kinglets, and hermit thrushes showed a preference for preburn 

conditions. Present as residents or peripheral visitors were juncos (Junco spp.), 

western tanagers (Piranga ludoviciana), and American robins (Turdus migratorius). 

Hansen (1978) states that Townsend’s warbler had an “almost invariable association 

with subalpine fir trees, which achieve their greatest density and foliage height 

diversity in this vegetation type.” He further states, “the community architecture 

contains a very large surface area of needles in which these birds forage.” 

In the control area, species composition in 1976 was similar to that in 1975 with only 

one additional species—a peripheral visiting red-breasted nuthatch (table 4). 

After fire, the bird community was radically different from that before burning. After 

fire, breeding bird species increased from 8 to 13. Five of these postburn species 

had not been recorded previously in the snag-thicket area (table 5). Four of the new 

species were single pairs of woodpeckers, including the hairy (Picoides villosus), 

common flicker (Colaptes auratus), black-backed (Picoides arcticus), and three-toed 

(Picoides tridactylus) (Hanson 1978). They nested in the charred, limbless spires of 

the snag-thicket burn area. According to Hanson (1978), this corresponds with 

Blackford’s (1955) observations that woodpeckers concentrate on burned forest. 

Hanson (1978) also observed that common flickers, black-backed woodpeckers, and 

Williamson’s sapsuckers (Sphyrapicus thyroides) were present at the still-smoldering 

site 3 days after the burn in 1975. Woodpeckers drilled holes in snags, which provided 

nesting sites for three pairs of mountain bluebirds (Sialia currucoides) and mountain 

chickadees during 1976 (table 5). Also in 1976, Hansen (1978) found increased 

numbers of aerial- and needle-feeding yellow-rumped warblers, and ground-foraging 

American robins and juncos in the burned area. In an unburned portion of the 

burned area, he observed a pair of bark-gleaning brown creepers (Certhia familiaris). 

Numbers of needle-foraging species, including Townsend’s warblers and hermit 

thrushes, declined precipitously after fire. Golden-crowned kinglets and western 

tanagers were absent from the snag-thicket burn area in 1976. 

Hanson (1978) suggests that the most significant finding of the bird population portion 

of the study was the change in diversity that occurred as a consequence of fire. In 

the snag-thicket burn area, species diversity increased significantly (P < 0.05) after 

fire. There was no change in the diversity index in the unburned control area 

between 1975 and 1976. Other bird population variables that increased after fire in 

the snag-thicket burn area were species richness and average number of species 

15 
Table 4—Total numbers and densities of breeding birds on the 12.5-hectare snag-thicket control area at 

the Table Mountain prescribed burn project in central Washington by year and foraging guilda 

1975 1976 

Species Guildb Pairs Species Guild Pairs 

Brown creeper BG 2.0 Red-breasted nuthatch BG .5 

Hermit thrush GF 2.0 Brown creeper BG .5 

Dark-eyed junco GF 2.0 Hermit thrush GF 1.5 

Mountain chickadee NF 1.0 Dark-eyed junco GF 1.0 

Golden-crowned kinglet NF 1.0 Mountain chickadee NF 1.5 

Yellow-rumped warbler NF,A 1.5 Golden-crowned kinglet NF 2.0 

Townsend’s warbler NF 5.5 Yellow-rumped warbler NF,A 1.0 

Townsend’s warbler NF 4.5 

Total 15.0 

Pairs/10 hectares 12.0 Total 12.5 

Pairs/10 hectares 10.0 

a After Hanson (1978).
 b BG = bark gleaning; GF = ground foraging; NF = needle foraging; A = aerial flycatching.


per census. Concurrently, these values declined in the control area between 1975 

and 1976. Refer to Hanson (1978) for detailed descriptions and maps of bird territories. 

Small mammals—In 1975, Hanson (1978) sampled small mammals by live-trapping 

on a 36-point grid with 15 m between points and two traps per point. Each grid was 

trapped for 7 days during the last week in July, August, and September 1975. In 

1976, the grid was expanded to 144 points. The same number of traps was used, 

but traps were rotated so that after 8 days, all 144 points had received four trap-nights 

of coverage. Each individual mammal was sexed, tagged, and its position noted in 

the grid. If the animal was recaptured, its identifying number and trap site were 

noted before release. Hanson (1978) made population estimates from multiple 

recapture frequencies using the modified Schnable technique (Overton 1965). 

Hanson (1978) was unable to determine home ranges for the mammals he trapped 

because of limitations of the study design. Small mammal population estimates 

were highly variable, partly because of the change in sampling techniques between 

1975 and 1976 (Hanson 1978). Also yellow pine chipmunk numbers appear to have 

declined between 1975 and 1976 for burned and control areas. Hanson concluded 

that the response of yellow pine chipmunk to burning was neutral or slightly positive. 

Hanson’s (1978) results also suggest that the number of Townsend’s chipmunks 

was severely affected by burning. The species was present in the snag-thicket and 

control areas in equal numbers before burning but absent from the snag-thicket 

burn area in 1976. Results were not definitive regarding the effects of burning, on 

any other species. Refer to Hanson (1978) for further details about the numbers of 

individual animal species found during trapping sessions and years for the snagthicket and control areas. 

16

Table 5—Total numbers and densities of breeding birds on the 12.2-hectare snag-thicket burn area 

at the Table Mountain prescribed burn project in central Washington by year and foraging guilda 

1975 1976 

Species Guildb Pairs Species Guild Pairs 

Red-beasted nuthatch 


American robin 


Hermit thrush 


Dark-eyed junco 


Golden-crowned kinglet 


Yellow-rumped warbler 


Townsend’s warbler 


Western tanager 


Total 

Pairs/10 hectares 

BG 2.0 

GF .5 

GF 4.0 

GF 1.5 

NF 3.0 

NF,A 2.5 

NF 14.0 

NF 1.0 

28.5 

23.0 

Red-breasted nuthatch 


Brown creeper 


Hairy woodpecker 


Black-backed woodpecker 


Three-toed woodpecker 


Common flicker 


American robin 


Hermit thrush 


Mountain bluebird 


Dark-eyed junco 


Mountain chickadee 


Yellow-rumped warbler 


Townsend’s warbler 


Total 

Pairs/10 hectares 

BG 1.5 

BG 1.0 

BD 1.0 

BD 1.0 

BD 1.0 

GF 1.0 

GF 2.0 

GF 2.5 

GF,A 3.0 

GF 3.0 

NF 1.0 

NF,A 3.5 

NF 2.0 

23.5 

19.0 

a After Hansen (1978).
 b BG = bark gleaning; GF = ground foraging; BD = bark drilling; NF = needle foraging; 


A = aerial flycatching. 

Forage Production, 

Forage Quality, Elk 

Forage Use, and 

Elk Diets 

Boltz (1979) studied the effect of the burn on forage production and quality, elk forage 

use, and elk diet preferences. Because elk are wide-ranging herbivores, Boltz 

(1979) assessed the importance of the forage produced and used on the prescribed 

burn site in the context of that of surrounding habitats used by elk. His studies 

included wet meadow, dry meadow, spruce swamp, open pinegrass-sedge, closed 

pinegrass-sedge, lupine-whortleberry, whortleberry, whortleberry-forb, sagebrush, 

clearcut areas, and the snag-thicket burn area (table 6). For each plant community 

type shown in table 6, Boltz (1979) determined the understory standing crop of forage 

by species, forage utilization by species, crude protein content by species, elk use 

and forage utilization, and elk food habits for each of these habitats. In this paper, 

only results of the various species groups, such as shrubs, grasses and grasslike 

species, and forbs, are presented. Refer to Boltz (1979) for detailed species 

information. Lupine-whortleberry with an overstory dominated by lodgepole pine 

and subalpine fir as a codominant comprised the largest proportion of total area 

(38 percent) (Boltz 1979) studied (table 6). The smallest proportion (0.9 percent) 

was occupied by the snag-thicket burn area. The sample period on which his results 

are based was June through September 1977. Whortleberry/forb lupine/whortleberry, 

and whortleberry habitats studied by Boltz (1979) are probably comparable to the 

snag-thicket burn area. At the burn area, he did not distinguish the snag area from 

the thicket area. 

17

Table 6—Elk habitats sampled in the area of the Table Mountain prescribed burn project in central 

Washington, including the snag-thicket burn areaa 

Percentage 

Community Overstory of total area 

Wet meadow None 2 

Dry meadow None 1 

Fir-spruce swamp Subalpine fir 6 

Open pinegrass-sedge Lodgepole pine-subalpine fir 4 

Closed pinegrass-sedge Lodgepole pine-subalpine fir 11 

Lupine-whortleberry Lodgepole pine-subalpine fir 38 

Whortleberry Lodgepole pine-subalpine fir 12 

Whortleberry-forb Subalpine fir-lodgepole pine 18 

Sagebrush None 1 

Clearcut None 6 

Snag-thicket burn area Lodgepole pine-subalpine fir 1 

a From Boltz (1979). 

Productivity of forage suitable for use by elk in summer 1977 was highest for 

the wet meadow (1379 kg/ha) and lowest for the burn area (34 kg/ha) (table 7). 

Productivity in areas comparable to the preburn setting (whortleberry-forb, whortleberry, 

and lupine-whortleberry) for the snag-thicket burn area ranged from 37 to 375 kg/ha 

(table 7). 

Boltz (1979) compared crude protein concentration (as a percentage of foliage dry 

weight) for 11 species found in the snag-thicket burn area with the same species on 

unburned sites monthly from June through September 1977. Burning enhanced 

crude protein concentration for every month of measurement. In June 1977, crude 

protein concentration for the 11 species averaged 23 percent for the burned area 

compared to 19 percent for the unburned area. Among species in the burned area, 

concentration ranged from 7 percent for whortleberry to 33 percent for lupine. 

Between June and September, average crude protein declined as foliage matured, 

but the difference between burned and unburned areas in September was about the 

same as in June (15 percent for burned compared to 10 percent for unburned). In 

September 1977 on the burned site, lupine was still highest and whortleberry still 

lowest in crude protein of the 11 species. 

Crude protein production (average of June through September 1977 values of Boltz 

[1979]) on a per-hectare basis was lowest (5 kg/ha) in the snag-thicket burn area 

(table 7). Commensurate with highest productivity, crude protein production was 

highest in the wet meadow. In the whortleberry-forb habitat, crude protein production 

was comparable to that of the snag-thicket burn area. It ranged up to 73 kg/ha in 

the lupine-whortleberry habitat exceeded only by wet meadow at 200 kg/ha. 

18

Table 7—Forage production, crude protein production, and elk use for habitats used by the Naneum 

Creek elk herd, including the snag-thicket burn area in summer 1977 at the Table Mountain prescribed 

burn project in central Washingtona 

Forage Crude protein 

Habitat productionb productionb Elk useb 

Wet meadow 


Dry meadow 


Fir-spruce swamp 


Open pinegrass-sedge 


Closed pinegrass-sedge 


Lupine-whortleberry 


Whortleberry 


Whortleberry-forb 


Sagebrush 


Clearcut 


Snag-thicket burn 


Kilograms per hectare Percent 

1,379 200 41.2 

323 37 7.7 

86 13 5.8 

388 41 13.4 

435 65 6.8 

375 73 9.5 

234 29 5.7 

37 6 2.0 

431 46 6.4 

427 39 Not included 

34 5 1.6 

a From Boltz (1979).


b Values averaged for June, July, August, and September sample dates.


Among all habitat types (excluding clearcuts), forage use by elk was greatest for the 

wet meadow and least for the snag-thicket burn area (table 7). Elk use of forage 

was higher in forested habitats comparable to the preburn conditions of the snagthicket area than in the snag-thicket burn area (table 7). 

Within each habitat type, elk diets were further characterized by percentage coming 

from each plant group: shrubs, grasses and grasslike species, and forbs (Boltz 1979) 

(table 8). For the June through July 1977 measurement, Boltz (1979) found that 

forbs constituted 90 percent of the diet for elk in the snag-thicket burn area. The 

primary forbs used were arnica, aster, and dragonhead. Elk sedge was the most 

heavily used of grass and grasslike species. Forbs were also a dominant part of the 

elk diet in lupine-whortleberry and whortleberry-forb habitats (81 and 86 percent, 

respectively). For the same period in the whortleberry habitat, shrubs (mainly 

whortleberry) were the primary forage used by elk. By September 1977 in the snagthicket burn area, percentage of shrubs and grasses increased in elk diets to 15 and 

27 percent, respectively. The proportion of forbs in the diet declined to 57 percent. 

Forbs remained the largest part of elk diets in September 1977 in lupine-whortleberry, 

whortleberry-forb, and whortleberry habitats (69 to 95 percent). 

19

Table 8—Percentage of elk diet from each class of plant in each habitat, including the snag-thicket burn 

area for the Naneum Creek elk herd in summer 1977 at the Table Mountain prescribed burn project in 

central Washingtona 

Sample period 

Habitat June-July August September 

Wet meadow 

Dry meadow 

Fir-spruce swamp 

Open pinegrass-sedge 

Closed pinegrass-sedge 

Lupine-whortleberry 

Whortleberry 

Whortleberry-forb 

Sagebrush 

Snag-thicket burn 

Percent 

Sb 0 0 0 

G 54.8 89.5 80.1 

F 45.2 10.8 20.0 

S 0 0 0 

G 26.3 76.6 100.1 

F 73.8 23.3 0 

S 0 10.4 0 

G 0 0 0 

F 100 89.6 100.1 

S 7.1 4.7 0.1 

G 71.4 62.5 99.8 

F 21.5 32.9 0 

S 10.8 10.9 0.1 

G 56.6 59.2 66.2 

F 32.6 29.9 33.8 

S 5.7 5.6 2.3 

G 13.7 18.7 13.1 

F 80.7 75.9 84.7 

S 61.5 21.5 13.2 

G 18.7 16.5 17.6 

F 19.7 61.9 69.2 

S 13.8 3.7 5.1 

G 0 0 0 

F 85.7 96.5 94.9 

S 0 4.4 24.4 

G 44.8 51.7 51.0 

F 55.2 44.0 24.7 

S 1.2 1.3 15.3 

G 8.7 2.0 27.3 

F 90.0 96.7 57.4 

a From Boltz (1979).
 b S = shrubs G = grasses/grasslike F = forb.


20

Conclusions Objectives of the fire prescription generally were achieved in this high-elevation 

subalpine fir/lodgepole pine habitat. In the snag burn area, the crown fire killed all 

the trees and removed most of the foliage. Overstory canopy cover was reduced by 

63 percent, and fuel loading was reduced an average of 71 percent (Woodard 1977). 

The duff layer (F and H or Oe and Oa) and fuels from 0 to 7.6 cm in diameter were 

reduced by more than 93 percent in the snag burn area. Fuels larger than 7.6 cm in 

diameter were reduced by 45 percent. In the thicket burn area, burning was spotty 

and the fire did not crown out. Overstory canopy cover was reduced by 18 percent. 

The lower fire intensity resulted because fuel accumulations were not as great as in 

the snag burn area. Fuel reductions reflected the lower fire severity in the thicket 

burn area. Average fuel reduction was 52 percent. Duff, fuels 0 to 7.6 cm in diameter, 

and fuels larger than 7.6 cm in diameter were reduced by 70, 34, and 34 percent, 

respectively. 

Woodard (1977) showed that as fire energy outputs increased, postburn plant cover 

decreased. If heat output exceeded 100 000 kcal/m2

, there was no reestablishment 

of the original plant cover. Woodard (1977, 1993b) also determined that plant 

diversity was reduced in both snag and thicket areas 1 year after burning, and 

that when heat output increased, species diversity declined. The thicket burn area 

exhibited lower species diversity than the snag area but had higher stability or 

resiliency than the snag burn area. Woodard (1977, 1993b) concluded that the 

preburn plant community composition was not a good predictor of the immediate 

postburn plant community. 

Woodard’s (1977) observations of low plant survival after fire suggest that most of 

the plants were not fire-resistant or did not have adaptive mechanisms that enabled 

them to survive a high-severity fire such as the one applied in this study. His work 

suggests development of the postfire plant community occurred by three principal 

mechanisms: (1) emergence of seedlings from species of heat-resistant seeds or 

those that remain viable for many years in the forest floor (dragonhead, arnica, and 

moss); (2) import of light-seeded species, such as subalpine fir, arnica, and sedges; 

and (3) seeds or rhizomes on microsite refugia that were sheltered from intense fire. 

Refugia were beneath large logs, at the downhill base of large-diameter trees, or 

near large rocks where fuel accumulations were low. Seedlings of heat-resistant 

plants contributed substantially to the first year cover after fire. Woodard (1977, 

1993a) concluded that the most important mechanism of regeneration is import from 

the perimeter of the burn and microsites in the burn interior that provide refugia for 

viable seeds of indigenous plants. He further suggested that the gene pool of interior 

microsites is of special significance for regeneration of large burn areas. 

Woodard’s (1977, 1993b) historical review of the fire history of the area and tree 

regeneration patterns suggests that the stand will replace itself, but that such a 

process will take a long time. 

Fire markedly altered several of the physical and microbiological properties and 

characteristics in the FL (F and H or Oe and Oa layers) and the upper 10 cm of mineral soil in the snag and thicket burn areas (Husby 1978). In the FL, pH was elevated by 1.9 to 2.2 units during the first 2 years after burning. Soil pH increased by 0.4 

to 0.7 units. Moisture contents of the FL and SL increased after burning in the thicket 

area only. Surprisingly, the fire had no detectable effect on mean temperatures of 

21

the FL or SL. There were, however, four sample dates in 1976 when the temperature 

of the FL of the snag burn area was 10 to 19 °C greater than that in the snag area 

control. Increased pH in the FL and SL provided an improved environment for bacterial growth that was reflected in the large increase in total bacteria in both burned 

areas in 1976 and 1977 (Husby 1978). Improved moisture in the thicket area likely 

facilitated the development of bacteria, but lower moisture levels in the snag burn 

area probably exerted a moderating effect on bacterial development. Bacterial 

numbers likely would have been higher in the snag burn area if moisture levels had 

been greater. Along with increased counts of total bacteria, proteolytic bacterial 

activity also increased (Husby 1978). Development of these organisms was facilitated 

by a higher amount of protein from the increased microbial pool and the postburn 

rise in N-fixing organisms. Increased numbers of bacteria were, however, not 

accompanied by an increase in microbial respiration from the FL. Microbial respiration declined significantly in the FL after burning. Husby (1978) indicated that Viro 

(1974) attributed this decline in respiration to the loss of the most easily used energy 

source for micro-organisms, the organic carbon of the forest floor. Nitrification and N 

fixation both increased after fire. Because N was likely limiting after the large losses 

from burning, increased N fixation would partially offset that limitation. Increased 

levels of nitrate after fire coincided with results of many other studies (Klemmedson 

and Tiedemann 1995). 

Fire resulted in an increase in the numbers of birds that established territories and 

nested in the area (Hanson 1978). New birds included four varieties of woodpeckers 

and western bluebirds. Diversity indices did not differ between 1975 and 1976 for 

the control area but increased significantly in the burned area between preburn and 

postburn years (Hanson 1978). Needle-foraging species declined after burning. 

There was a sharp decline in numbers of Townsend’s warbler and hermit thrush, 

and disappearance of golden-crowned kinglet and western tanager. 

Hanson’s (1978) study showed that numbers of Townsend’s chipmunk declined 

sharply following burning. Yellow pine chipmunk was the only species of the five 

sampled that showed a neutral or positive response to burning. 

Heat output of the fire, particularly in the snag area, was sufficient to severely limit 

development of vegetation the first 2 years after burning (Woodard 1977, Boltz, 

1979). This area had the lowest usable forage for elk of any of the habitats that 

Boltz (1979) sampled in summer 1977. Forage quality, as indicated by increased 

crude protein, was improved by burning, but low productivity and distance from 

water reduced the value of the area for elk forage. 

Results of these studies suggest that, in the short term (2 years), the fire exerted a 

detrimental effect on the forest floor and understory vegetation. The potential benefits of releasing the understory vegetation to provide forage for the resident elk herd 

were not realized because of the poor development of vegetation. Nearly complete 

elimination of the duff and litter on the forest floor in the snag area and in some 

parts of the thicket area was likely associated with substantial losses of essential 

plant nutrients, such as N (Raison and others 1984), S (Tiedemann 1987), and 

phosphorus (P) (DeBano 1991, Raison and others 1984), especially at the temperatures achieved during this fire. 

22

The short-term period of postfire studies limits the application of the information. 

However, the manner in which the study was designed and set up in a remote area 

and the extensive baseline data provide an unprecedented opportunity to explore 

longer term (24 years) effects of fire on this forested ecosystem. It would seem 

appropriate to now conduct studies that examine the same parameters. 

Acknowledgments	 The authors appreciate the thorough, candid technical reviews provided by John 

Lehmkuhl, PNW Research Station, Wenatchee, WA, and Marvin Hoover, Rocky 

Mountain Research Station (retired), Wenatchee, WA. 

English 

Equivalents 

Literature Cited 

When you know: Multiply by: To find: 

Centimeters (cm) 0.39 Inches 


Meters (m) 3.28 Feet 


Square meters (m2

) 1.20 Square yards


Kilometers (km) 0.62 Miles


Hectares (ha) 2.47 Acres


Square meters per hectare (m2/ha) 4.36 Square feet per acre 


Metric tons per hectare (t/ha) .445 Tons per acre


Kilocalories per square meter (kcal/m2) .369 BTU per square foot 


Milligram (mg) .000035 Ounces 


Micrograms per gram (μg/g) .000001 Ounces per ounce


Kilograms (kg) 2.205 Pounds


Kilograms per hectare (kg/ha) .89 Pounds per acre


Milliliters (mL) .034 Fluid ounces 


Celsius (°C) 1.8 and add 32 Fahrenheit 


Alexander, M. 1961. Introduction to soil microbiology. New York: John Wiley and 


Sons, Inc. 472 p.


Binkley, D. 1986. Forest nutrition management. New York: John Wiley and Sons,


Inc. 290 p.


Blackford, J.L. 1955. Woodpecker concentration in burned forest. Condor. 53: 28-30.


Boltz, M.J. 1979. Impacts of prescribed burns and clearcuts upon summer elk food 


habits, diet quality, and distribution in central Washington. Pullman, WA: 


Washington State University. 129 p. M.S. thesis.


Brown, J.K. 1974. Handbook for inventorying downed woody vegetation. Gen. 


Tech. Rep. INT-16. Ogden, UT: U.S. Department of Agriculture, Forest Service,


Intermountain Research Station. 24 p. 


Byram, G.M. 1958. Combustion of forest fuels. In: Davis, K.P., ed. Forest fire: 


control and use. New York: McGraw Hill: 61-123.


23

DeBano, L.F. 1991. The effects of fire on soil properties. In: Harvey, A.E.; 

Neuenschwander, L.F., comp. Proceedings, management and productivity of 

western-montane forest soils. Boise, ID. Gen. Tech. Rep. INT-280. Ogden, UT: 

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151-156. 

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ecosystem health assessment. Vol. 1: Executive summary. Portland, OR: U.S. 

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burning and select tree removal (Grass Camp, 1975). Res. Pap. PNW-234. 

Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest 

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Franklin, J.F.; Dyrness, C.T. 1988. Natural vegetation of Oregon and Washington. 

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Intermountain Research Station. 86 p. 

Hoover, M.D.; Lunt, H.A. 1952. A key for the classification of forest humus types. 

Soil Science Society of America Proceedings. 16: 368-370. 

Hu, L.; Youngberg, C.T.; Gilmour, C.M. 1972. Readily oxidizable carbon: an index 

of decomposition and humification of forest litter. Soil Science Society of America 

Proceedings. 36: 959-961. 

Husby, G. 1978. Survey of microbiological activities in a pine forest soil following a 

prescribed burn. Ellensburg, WA: Central Washington University. 81 p. M.S. thesis. 

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Klemmedson, J.O.; Tiedemann, A.R. 1995. Effects of nutrient stress. In: Bedunah, 

D.J.; Sosebee, R.E., eds. Wildland plants—physiological ecology and developmental 

morphology. Denver, CO: Society for Range Management: 414-439. 

McColley, P.D. 1976. Soil resource inventory, Wenatchee National Forest. 

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Northwest Region, Wenatchee National Forest. 318 p. 

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Overton, W.S. 1965. A modification of the Schnable estimator to account for the 

removal of animals from the population. Journal of Wildlife Management. 

29: 392-395. 

Page-Dumroese, D.; Harvey, A.; Jurgensen, M.; Graham, R. 1991. Organic 

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Neuenschwander, L.F., comp. Proceedings, management and productivity of 

western-montane forest soils. Gen. Tech. Rep. INT-280. Boise, ID: U.S. 

Department of Agriculture, Forest Service, Intermountain Research Station: 95-100. 

Raison, R.J.; Khanna, P.K.; Woods, P.V. 1984. Mechanisms of element transfer to 

the atmosphere during vegetation fires. Canadian Journal of Forest Research. 

15: 132-140. 

Tiedemann, A.R. 1987. Combustion losses of sulfur from forest foliage and litter. 

Forest Science. 33: 216-223. 

Tiedemann, A.R.; Klock, G.O. 1977. Meeks Table Research Natural Area: 

reference sampling and habitat classification. Res. Pap. PNW-223. Portland, OR: 

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watershed nutrient economy following wildfire and fertilization in eastern 

Washington. Journal of Environmental Quality. 7: 580-588. 

Tiedemann, A.R.; Klemmedson, J.O.; Bull, E.L. 2000. Solution of forest health 

problems with prescribed fire: are forest productivity and wildlife at risk? 

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and ecosystems. New York: Academic Press: 7-45. 

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Harvey

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25

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26

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