i This· fi About e was ThI.s F created ile· •nv,t l.ssc . bY sca a ns 'd n ni e n ' ntified g the ho.!'\fe Pr.mt ed bY t he ver, s softw PUb/'lc o a ------------...:.·..:::... rne rnls at r e ha ve takes been tnay re correc main . · . .. 6 Effects of Slash Burning on Plant Succession and Timber Stand Establishment R. E. Bigley and J. A. Henderson ABSTRACI' Conifer establishment and growth on burned and unburned areas vary · considerably by site. This paper deals with the influence of bum severity on the environment, vegetation response after burning, conifer establishment and early growth, and documentation of variations in effects of burning on plant succession. Controlled burning of logging slash in the Pacific Northwest strongly influences the initial environ­ mental conditions in which vegetation must develop and the possible plants that may establish. Knowledge of the subject is limited both by the few sites where direct comparisons of burned and unburned plots have been made and the short duration of most observations, but enough is known about several widespread species to predict their response to prescribed f1re. The literature suggests that conifers planted on burned areas in coastal Washington and Oregon gain an advantage from reductions in competing vegetation but may suffer from increased animal damage. The conclusion is reached that information on the effects of burning on vegetation could be better utilized if respose data were stratified by forest zones and plant associations (or groups of associations). Controlled burning of logging slash is a major influence on initial plant succes­ sion and early stand development of Pacific Northwest forests. The extent of our knowledge, however, is limited both by the relatively few sites where direct comparisons have been made on paired burned and unburned areas arid the shon duration of most observations. Enough is known about the response of some species to prescribed frre to give general predictions of their response. Variation among sites and variation in the influence of burning on vegetation and the environment will continue to complicate the transfer of information about slash burning effects to potential users. Succession is not only a sequence, as described by a series of plant species and abundances, but a process in which the environment and thus ihe suitability of the site for different species change. The main effect of slash burning is to modify the environment and influence species survival, which in tum influences processes such as initial plant establishment and growth. The effect is ultimately reflected in differences in vegetation on slash burned and unburned areas. This Bigley and Henderson paper focuses on those changes that take place in the first two decades of stand growth, with emphasis on areas west of the Cascade crest in the Pacific Northwest This period includes the most rapid changes in vegetation and typically the establishment of a conifer canopy. Once closed, a canopy has considerable influence on slowing the rate of vegetation change. We will fmt examine the major effects that burning has on the environment in which vegetation develops. Then we will discuss the general state of knowledge about burning effects on herbaceous and woody plants, and, more specifically, conifer regeneration and early growth. We will conclude with a few ideas on how to utilize existing information dealing with burning effects on succession. Influence of Burn Severity on the Environment The term "burn severity" as used in this paper refers to the postfll'e surface conditions as described by Morris (1958). A burn of low severity would leave the forest floor intact, consuming only the litter layer. A severe burn would remove the forest floor, and usually discolor the surface of the mineral soil. Slash burning rarely affects an area evenly. Describing a site as "burned" may give little information as to the extent of impact on site or vegetation. The variable influence of slash burning has made the detailed interpretation of treatments difficult over entire slash burned units. The severity and distribution of burning effects are related to the moisture content, amount, and distribution of fuels, but can vary with ignition patterns, topography, and weather conditions. Spring bums are generally lower in severity because of higher fuel moistures and result in less reduction of forest floor and exposure of mineral soil than summer and autumn burns (Hawkes 1986, Little et al. 1986). In studying the effects of slash burning on fire hazard and revegetation, Morris (1958, 1970) conducted extensive line intersect sampling of ground conditions after broadcast slash burning. These records allowed the reconstruc­ tion of the distribution of burn severity over a 0.25 acre plot shown in Figure 1. Intervals between line intercept transects are averaged, resulting in an over­ simplification of the actual pattern. The pattern and size of affected areas vary considerably over the plot, creating a wide range of microsites for new plant establishment Where unburned patches of vegetation are adjacent to moderate­ ly or severely burned areas, existing shrub canopies or below-ground growth could easily expand and occupy the burned space. Both the size of a burned area and the condition of the neighboring area may influence the type and rate of regrowth at a given location. Stratification of burning effects by severity classes provides good informa­ tion for specific small locations, but little guidance on stand-level effects. This is because of the tremendous variation possible in the proportion and distribution of areas of different bum severity and the response at different sites. Several studies provide examples of how plant response differs in areas of different bum Effects on Plant Succession .Severe Figure 1. .Light .Moderate Bum severity distribution on a 0.25 acre Unburned plot, broadcast-burned area near Lowell, Oregon. severity. Morgan and Neuenschwander (1988) reported differences in shrub cover response on 172 square foot plots having at least 90% of bums of low or high severity on a speCific site type in Idaho. They concluded that shrub community differences following low and severe bums were greatest in the first three years after burning; thereafter, composition became more similar with time. Regrowth from species with or without rhizomes was greater on the low than on the high severity bum locations, but shrub seedlings were more prolific on areas of severe bum. Both the composition and frequency of occurrence of species differed between the areas of bum severity. Halpern (1987) found that the duration of herb and shrub dominance differed with site type and bum severity. On one site, the duration of the herb layer dominance decreased with increasing bum severity because both redstem (Ceanothus sanguineus) and evergreen ceanothus (Ceanothus velutinus) showed greater development on burned sites and shaded the herbaceous plants. Slash burning has many possible effects on the characteristics of the soil, ecosystem processes, and the microclimate of a site which may also influence the vegetation that reestablishes. Several works deal with these topics in detail (Neal et al. 1965, Feller1982, Ahlgren and Ahlgren 1960, Boyer and Dell1980, Barnett 1984, de Montigny and Auclair 1982, Grier 1972). Changes to the soil surface are particularly important to plant establishment and initial growth on burned sites. To some extent burning increases overland flow of surface water by the removal of forest floor and live vegetation, and increases the removal of seed and young plants by erosion. Drying of the soil surface as a result of the loss of organic matter and increasing surface tempera­ Bigley and Henderson tures in summer may prevent seed gennination and survival of seedlings. Boyer and Dell (1980) noted that soils under some plant associations are more prone to develop water repellent surfaces than others after burning because of volatilization of organic compounds during burning. Ambient air and soil surface temperatures generally display wider extremes following slash burning (Lafferty1980) by the reduction of thennal cover ( both forest floor and vegetation) and changes in surface albedo. In the winter, surface soil temperatures can be lower on burned than unburned sites, resulting in frost heaving of seedlings' roots or foliage damage on burned areas. Burning generally causes temporary increases in available nutrients. Isaac ( 1940) speculated that increased nitrogen availability on burned sites may result in development of conifer foliage in advance of roots, leaving them susceptible to drought; however, this increased nutrient availability is generally seen as a benefit to early growth of reestablishing vegetation. Failure to reestablish the next conifer crop promptly after burning could influence succession for a considerable time if nutrients are lost from the site or allocated to competing vegetation. Vegetation Response after Burning Modes of Plant Regrowth The most visible influence of burning on succession is the removal of part of the existing vegetation. Plants can survive burning both as seed and as aboveground and underground structures. Survival of established plants through a bum is related to the depth and type of root system (McLean 1969). Plants with shallow roots are susceptible even to light bums. Shrubs with rhizomes within the mineral soil are rarely seriously damaged by burning. Miller (1977) found that the number of blue huckleberry (Vaccinium globulare) sprouts was affected by the depth of heat penetration. Low severity spring burns resulted in multiple new shoots arising from a single old branch through stimulation of buds along the rhizomes. In this case, stem density could increase 80 to 120%. Autumn burns were more likely to reduce stem density because of greater heat penetration. The season of burning can be important for the vigor of regrowth, because carbohydrate reserves in underground tissues are low in the summer and highest in the autumn (Volland and Dell1981). It is well known that delayed seed gennination can result in large reserves of buried seed within the forest floor and mineral soil. Although the numbers of buried seed may vary greatly, the distribution of seed in an undisturbed profile decreases sharply with depth (Figure 2 ). The number of seeds and the species composition may vary with the longevity of the seed and resistance to frre. In some cases seeds can persist in the soil for many decades (Roberts1981). Pratt et al. (1984) studied a ponderosa pine community in eastern Washington and found viable seed of flfty-seven species in the surface soil layers, twenty- Effects on Plant Succession Seeds/tt2 0 40 20 LFH 0 .8 (/) Q) - ..c CJ c: 1.6 2.4 4.0 0 - ..c Q. Q) c Ceanothus ve/utinus 200 100 LFH 0 .8 1.6 2.4 Stellaria media 4.0 Figure 2.. Distribution and number of buried viable seed per square foot of two species. Source: Weatherspoon (1985), Pratt et al. (1984). one of which were not represented in the aboveground vegetation. In Oregon and Washington, Ceanothus species provide the best known example of burning as a stimulation to germinate seed that has been buried for long periods (Gratkowski 1961). Orme and Leege (1976) found that seedling emergence, survival, and growth ofredstem ceanothus (C. sanguineus) was greater after fall than spring bums. Burning can remove some buried seed by heating to lethal temperatures. Below that zone, buried seed may benefit from heat scarification; seed that lies still deeper may remain viable and dormant. Recruitment of new seedling s may also be influenced by the remaining aboveground vegetation. Shading can inhibit germination of most species, but to a lesser extent for species charac­ teristic of later successional stages (Pratt et al. 1984). Determinants of Vegetation Development The course of succession after a disturbance such as slash burning has two general detenninants: the potential plant species available and the specific environment. both of which include chance. The species composition of the site before burning exerts a strong influence on subsequent vegetation development (Dymess 1973; Halpern, in press; Bigley, in preparatioQ). Besides the initial aboveground vegetation, the influx of seed is very important, especially on burned sites. Many wind-dispersed weedy species are typically found on burned sites. Kellman (1974) found seed input for the fli'St three years after clearcutting on a site in coastal British Columbia to be 60, 55, and 23 per square foot, and Henderson respectively, for years one, two, and three. Vegetation on unburned sites is usually composed of a greater proportion of remnant and persistent vegetation. Environmental characteristics of the site, including historic factors such as burning impacts, comprise the second determinant of succession. The specific characteristics of the site can affect the rate of succession and prolong or reduce the influence of burning effects (Bigley, in preparation). At high elevations where colonization and growth rates are slower, even low severity bums can reduce plant cover for decades, bycreating poor conditions for reestablishment. Because of the potential for many plant species tocontributeto a community, and the various impacts of burning combined with chance occurrences, it is not surprising that plant succession can take different paths on burned sites com­ pared with unburned ones (Connell and Slatyer 1977; Cattelino et al. 1979; Halpern, accepted; Bigley, in preparation). Amo et al. (1985) have provided a good example of how chance history of a site and burning interact to influence early plant succession (Figure 3). In this example, representations of plant T S R UCT URAL NO. Com­ mun­ ity 1 SH RU · B HER B SAPUNG STA GE S POLE MATURE FORE T S OD L GROWTH FORE S T VAGL A V GL �������rr� types U bu n r e nd u Br e nd B ur n ed with C EVEseeds Figure 3. Idealized multiple pathway of succession in a Douglas-fir (Pseudotsuga menziesii, PSME) and blue huckleberry community (Vaccinium globulare, VAGL) showing the effects of burning depending on stand history. Species abbreviations: evergreen ceanothus (Ceanothus velutinus, CEVE), lodgepole pine (Pinus contorta, PICO), pinegrass (CalQITUlgrostisrubescens, CARU). Source: Amo et al.(1985). dominance are shown for unburned sites and for sites burned with and without ceanothus seed in the soil. Removal of the shrub community with severe burning (communities 2 and 3) resulted in two possible paths· of community develop­ ment, depending on the presence of ceanothus seed. Effects on Plant Succession Broadcast Burning Information on early succession after slash burning generally comes from short-term observation of often poorly characterized sites. Chronosequence approaches are good for deriving long-term trends; however, it is often impos­ sible to determine if differences in the treatments and conditions that occurred (1982) on the different sites gave rise to the communities observed. Franklin emphasized that there is no substitute for observation over time in the study of plant succession. Few studies have reported observations from matched burned and unburned areas for an extended period. There is a scattered and disjointed data base for the response of vegetation to prescribed slash burning west of the Cascade crest in the Pacific Northwest (Table 1). Most reports consist of observations, general!y from three to ten years, and provide only general trends in plant cover. Vegetation development records from repeated sampling of permanent plots over twenty-five years come from only two plot networks. The most extensive are those established by Moms from 1946 to 1951 and consist of roughly 0.25 acre plots in burned and unburned pairs originally at sixty-two locations in Washington just north of Mount Rainier to south of Roseburg, Oregon. As of 1988, forty-nine pairs remained intact for continued sampling. The second series of small permanent succession plots, with observations through twenty-five years, was established by C. T. Dyrness in 1962 on two experimental watersheds in the H. J. Andrews Experimental Forest, Oregon. A common feature of studies listed in Table 1 is combination of data from sites that are not similar. This practice results in large variations and the inability to compare results with studies having a different combination of sites. To illustrate how different some plant responses can be on different sites, Figure 4 shows plant cover on two plant associations on the Lowell area of Oregon. Both areas burned at similar severities, but species response was different on the two plant associations. On the western hemlock and sword fern site, thimbleberry increased on the burned plot, probably as a response to growing space made available by the removal of other species such as vine maple, mineral soil exposed by burning, and perhaps the increase of available nutrients; salal was completely removed from the burned area for many years after the frre. On the rhododendron site, thimbleberry grew poorly on the burned plot, as did most species on this site (where summer moisture stress can be substantial). Rhododendron regrowth was delayed for several years by burning. Valuable information could be lost if the results from these two sites were combined. Since the studies in Table 1 cover a wide range of sites and bum severity, general conclusions cannot easily be made; however, a pattern in early succes­ sion is commonly reported which shows that both remnant vegetation and opportunistic invaders inhabit burned and unburned sites. Burning reduces the predisturbance flora early in the sere and shifts to a greater preponderance of invading species (e.g., West and Chilcote 1968). Shrub regrowth is usually slowed on burned areas compared with unbwned ones. Bigley and Henderson Table 1. Summary of srudies comparing vegetation on bmned and unburned areas in western Washington, Oregon. and British Columbia. Authors lngTam ( 1931) General Location Number of Study Areas Wind River Experimental Forest near Carson. Washington Years of Postbuming Observation 10 Reid et al. ( 1938) Eight observations over ten years on small replicated burned and unburned plots Shows effects of rebuming and grazing Added statistical analysis and discussion from Ingram 1931 Kienholz (1928) Southern Puget Sound basin, Washington Isaac Western Washington and Oregon (1940) Comments on Data Reported 7 3 Small plots stratified by bum severity, slope, and aspect IS 8 Permanent plots examined annually, not paired Data confounded by combining results from burned and unburned plots General trends of speci.fic species reponed Yerkes (1960) H. J. Andrews Experimental Forest, Oregon Feller UBC Research Forest near Maple Ridge, B.C. (pers. comm.) Morris (1958) Western Washington and Oregon 14 s 2 1 and 7 62 7 Areas ranging from 1,850 to 3,800 feet in elevation combined in different groups from year to year for average species occurrences Detailed data not presented for unburned plots Data available for individual species Good description of srudy areas Several observations of small plots on paired treatment areas Effects on Plant Succession Table 1. Continued. Authors General Location Number of Study Areas · Years of Postbuming Observation Comments on Data Reported 62 11-16 Morris 1958 updated Overall averages for Cascades and coastal areas presented Oakridge, Oregon 2 13 Photographs from plots established by Morris to year 13 Steen (1966) Oakridge, Oregon 13 11-16 More detailed treat­ ment of a subset of the plots treated by Morris (1970) Kraemer (1977) Western Washington and Oregon 33 25 Detailed update of some of the areas covered by Morris (1970) Individual species Morris (1970) Western Washington and Oregon Steen (1965) discussed Bigley Western Washington and Oregon Dymess (1965) H. J. Andrews (in prep.) 49 37-40 2 Comparison of pre- and postlogging and post­ bmning vegetation Experimental Forest, Oregon Dymess (1973) H. J. Andrews Halpern (1987) H. J. Andrews Halpern H. J. Andrews 2 5 Experimental Forest, Oregon Experimental Forest, Oregon (pers. comm.) Experimental Forest, Oregon Detailed update of some of the areas covered by Morris (1970) and Kraemer (1977) Focus on vegetation development between plant communities Annual observations on smJill pennanent plots Irtdividual species data by year 2 17-21 Update of Dymess (1973) Detailed analysis of bmn severity and plant communities Update of previous Bigley and Henderson 24 -.------. 12.0-r------, 21 Percent cover 9.0 18 Percent cover 15 12 9 T/lhnbleberry 6.0 3.0 Sala! Percent cover 13 Unburned 26 36 Burned Figure 4. Cover of selected species on burned and Wlbumed plots on two different plant associations: Left: western hemlock-sword fern association. Right: western hemlock­ rhododendron association. Conifer Establishment and Early Growth Advance Conifer Regeneration Advance regeneration makes a major contribution to stocking on unburned areas in many regions (Heavilin 1977, Isaac 1943) but is largely removed from burned areas. Natural conifer regeneration differs substantially by plant associa­ tion (Shearer 1985) and forest zone. The potential importance of advance regeneration depends on its abundance, species composition, and the prospects for successful artificial regeneration. The three major vegetation zones in the Cascades (Figure 5) show substantial differences in advance regeneration. The number of seedlings (between 6 inches in height and 1 inch dbh) was found to be 1,320, 24,834, and 43,380 per acre, respectively, for the western hemlock, silver fir, and mountain hemlock zones (Figure 6). The number and species composition of saplings in each zone reflect the potential crop composition. We are unaware of any data on the comparative survival of advance regeneration in the different forest zones of the· Cascades. Where artificial regeneration can be difficult (e.g., mountain hemlock zone and some areas of the silver fir zone), loss of advance regeneration can significantly reduce stocking for many years (Bigley and Miller, in preparation). On other sites, burning is an effective means to control species composition by the removal of advance regeneration. Effects on Plant Succession 6000 5000 Q) - Q) :!::. c: 0 ·.;: co > Q) [jj N s 4000 PACIFIC SILVER 3000 FIR ZONE 2000 WESTERN HEMLOCK ZONE 1000 0 Figure 5. Relative position of the major forest zones west of the Cascade crest in relation to aspect and elevation. 100000 WESTERN HEMLOCK SILVER FIR ZONE ZONE MTN HEMLOCK ZONE 10000 Q) ;;; a. VI Q) 1000 100 10 2 I 3 iI 4 5 2 3 4 I 5 2 3 4 5 Size Class E] PSME . TSHE m THPL ABAM 0 TSME Figure 6. Number and species distribution of tree saplings in the major forest zones. Abbreviations: Douglas-fir (Pseudotsuga menziesii, PSME), western hemlock (Tsuga heter ophylla, TSHE), western redcedar (l'huja plicata, THPL), Pacific silver fir (Abies amabilis, ABAM), mountain hemlock ([suga mertensiana, TSME). and Henderson Conifer Establishment and Stocking The feasibility of conifer regeneration within a reasonable time is a major factor in deciding whether to burn. Planting costs may not differ between burned and unburned sites if some yarding of unmerchantable materials is done (Lance Raff, USDA Forest Service, Darrington, pers. comm.). However, seedling distribution, planting quality, and early growth rates also must be considered. Much of the existing infonnation about conifer establishment on burned sites focuses on natural regeneration. Most dat3. on planted seedling success and growth rates on burned and unburned sites (such as stocking inventory results) never fmd their way to the published literature. The data that are published reflect the same variation and contradictions seen from site to site in understory succession resulting from differences in burn severity and the site environments. In the first half of this century, a considerable effort was made to determine the effects of slash burning on establishment of natural conifer regeneration. Conclusions consistently emphasized the importance of the timing of the burn and dispersal of the conifer seed crop (Allen 1946, Silen 1952). Research into specific conditions comparing germinant survival on burned and unburned sites has found the blackened surface of burned plots hazardous to germinants (Kienholz 1928, Hermann and Chilcote 1965). Lavender et al. (1956) and McCulloch ( 1944) emphasized a balance between moderate burn severity and slash loading to ensure regeneration success. In studies of burned areas, tree seedling survival, stocking, and growth results, data are often stratified by burn severity class or simply presented as an average response. It is difficult to compare the results of studies that report either the general or microsite specific responses and provide no way to estimate the other measure. Stratification of burned areas provides a good idea of the relative performance of seedlings on those specific sites. The conclusions of different studies on tree seedling response on burned and unburned areas are often contradictory. Several authors have found poor sur­ vival on areas exposed to extreme burning (Isaac and Hopkins 1 937, Austin and B aisinger 1955, Baker 1968). However, Miller and Breuer ( 1 984) reported that survival was the same on burned and unburned areas with little brush competi­ ings tion after three years. Burning can also increase the survival of planted s (Vyse and Muraro 1973). Each study needs to be examined to detennine where and under what situations the conclusions can be applied. Combining of data from many widely distributed s tes can provide very general but useful regional trends. Morris (1970) gave general trends in primari­ ly naturally regenerated conifer stocking on sixty-two paired burned and un­ burned plots. Burned plots in the Oregon Coast Range regenerated much quicker than plots on the west slope of the Cascades. Although by year 12 there were no differences between the burned and unburned coastal plots, the Cascade plots found signs of having greater stocking on the unburned plots. On closer analysis, Bigley (in preparation) found considerable differences in stocking rates between different plant associations on Morris's plots. Reports from unconfounded, paired, burned and unburned plot studies are 58 Effects on Plant Succession rare. Isaac (1943) found that regeneration ofDouglas-frr occurred faster and in gr ter numbers on unburned plots compared with paired burned plots where other conditions were comparable. Munger and Matthews (1941) presented results of stocking surveys at six burned and unburned locations on coastal Washington and Oregon. At the end of seven years the unburned areas had over five times as many seedlings as the burned areas. They also cited other data from the same area that suppon this trend, but data were taken from unpaired units. Gockerell (1966) reported results of a 4,130 acre survey (by the Washington State Department of Natural Resources) of planted clearcuts in northwestern Table 2. Survival and distribution of 1- to 4-year-old planted seedlings in burned and unburned areas of 4,130 acres of planted clearcuts, 1958-63 (Gockerell 1966). Unburned Areas Burned Areas Planted survival (%) 57 55 Planted stocking (%) 38 42 Planted and natural stocking(%) 79 69 Animal damage(%) 28 78 Planted Douglas-flr, average height(ft) 4.1 3.4 Washington around 1960. Survival and distribution of 1- to 4- year-old planted seedlings were about the same on the burned and unburned areas; however, if natural regeneration was included, stocking was greater on the unburned areas (Table 2). Although not from paired areas, this srudy is valuable because of the extensive area surveyed. Early Growth of Conifers on Slash Burns Little published information exists on direct comparisons of tree growth on matched burned and unburned areas (Table 3). Available data typically cover only the frrst few years of growth and may misrepresent the longer term growth trends. Beese (MacMillan Bloedel Ltd., pers. comm.) compared growth rates of planted Douglas-frr on burned and unburned areas and found the first year growth gave little indication of the second year results. Seedlings planted in intact forest floor grew best the frrst year, but seedlings on mineral soil humus grew better the second. Slash burning has a major influence on early growth by the reduction of competing vegetation, particularly shrubs. This reduction in competition may explain many of the differences observed in initial conifer growth on matched burned and unburned sites. Vihnanek and Ballard (1988) examined stocking, growth, and foliar nutrient levels of planted Douglas-frr on burned and un­ burned, salal dominated sites on the east side of Vancouver Island. Burning Bialey and Henderson Table 3. Summary of studies comparing tree growth on burned and lll1bu."11ed areas in western Washington, Oregon, British Columbia. and northern California. Number General Authors Years of Effect of of Study Postbuming Burning on Areas Location 2 Cascades Tarrant and Wright Observation 1 and 2 Growth None Oregon ( 1956) coast N aturally regenerated Douglas-fir height Root lengths presented (1955) Ruth Comments* 5 None Douglas-fir height growth Heavy brush competition on paired burned and unburned Madison (1959) 2 Oregon coast 8 None areas Sitka spruce height growth Compares sites with different aspects Knight ( 1961) Vancouver Island, B.C. 1 2-3 Negative Douglas-fir diameter and height growth Limited salal competition deMon- Vancouver tigny Island, B.C. 1 4 Negative Chronosequence study Douglas-fir height and (1985) growth de..'"Teased with increasing bum severiry Heavilin KlamathN.F., (1977) California 3 7 Negative Douglas-fir height growth Advance regeneration may have contributed to the taller trees on the unburned areas Gockerell ( 1966) Olympic Peninsula, Many 1-4 Negative 3 Positive Douglas-fir height growth Washington Miller and Oregon Breuer Douglas-fir height growth Survey results \ ( 1984) Stein Oregon (1986) coast 6 5 Positive Douglas-fir height growth Paired plots replicated at a location Vihnanek Vancouver and B allard Island, ( 1988) 20 5-15 Variable Douglas-fir height growth Each location with paired burned and unburned B.C. *If not specifically stated. trees were planted; although investigators mention difficulty in difierentiating between planted and naturally regenerated Douglas-fir if the areas had been planted more thm a few years. 60 Effects on Plant Succession 9A o��r-��� ���� 5 6 7 8 910 8 910111 2131415 ���--��� 8 91011 1 2 131415 Stand Age lyearsl Figure 7. Generalized relationships of Douglas-flr height growth on sites on Van­ couver Island. Source: Vihnanek and Ballard (1988). clearly reduced competing salal cover, increased Douglas-flr height and diameter growth, and improved the status of several foliar nutrients compared with conifers on the unburned portions in the 5- to 15-year-old stands measured. Average tree diameters on eighteen of the twenty sites were greater on burned plots. Height growth was characterized in three general curves (Figure 7). The fl.l'st curve (A) shows similar growth rates on burned and unburned areas; the others show greater growth on the burned areas. The extreme case (curve C) shows growth between burned and unburned conditions as increasingly diver­ gent. Although burning may result in initially superior growth, there are cases where growth trends on burned and unburned sites have crossed after seven to nine years (Braathe 1973). This is about the same time that rapid rates of mineralization promoted by burning would be expected to decline (de Montigny and Auclair 1982). In contrast, several studies have reported that initial height growth is greater on unburned sites than on burned ones. Height growth trends can be deceptive, however, because animal damage is usually greater on burned sites (e.g., Gockerell 1966, Stein 1986) and may be a source of error that discriminates against "growth" on burned sites in much of the literature. Stein (1986) found Table 4. Effect of use of vexar rubes on plant survival and growth on burned and un­ burned sites(Stein 1986). Height(ft) Survival(%) Use of Vexar Tubes Burned Unburned . Burned Unburned Us ed 88 83 5.6 5.0 Not used 59 46 4.6 3.8 Bigley and Henderson that protection from animals by using vexar tubes increased height growth of Douglas-fir on both burned and unburned sites (Table As with vegetation response to burning, tree 4). growth response varies from location to location and appears influenced by bum severity. Stem diameter growth close! y reflects the growth of the root system and thus the establishment of a seedling, although few studies report diameter. Few reports provide enough information on the study site environment, plant competition, and occurrence of animal damage to transfer results to other areas with assurity. Documenting Variations in Effects on Plant Succession Differences in site type (e.g., plant association and soil type) and severity of a bum strongly influence the establishment and growth of both conifers and nonconifer vegetation on burned areas. Many of the contradictions evident in the literature result from lack of recognition of the importance of these factors. We suggest that information on the effects of burning on vegetation could be better utilized if response data were stratified by forest zones and plant associa­ tions (or groups of associations). Considerable information exists in the form of observations and experience of local land managers, but is not available or is not being used by those who should have access to it The following is an idea how this valuable locally based information could perhaps be better utilized to support local problem solving Wildlife Re eneralion WET Burn as needed l£d Moisture Avoid hot burns Consider not burning 0 Figure 8. An idealized framework to compile and transmit information on burning effects on different forest ecosystems. Effects on Plant Succession and planning. A general framework to compile and ttansmit infonnation on burning effects is shown in Figure 8. Infonnation is grouped on a site-specific and objective-specific basis. To allow site specificity, observations are arran ged by forest zone and plant assoc iation. The ordination of plant communities in this example was generally based on a diagram of the western hemlock zone on the Gifford Pinchot National Forest (Topik et al. 1986). The areas of similar re­ sponse or recommendations are defined based on the expected need and effects that broadcast slash burning will have for a particular objective. In the example shown for vegetation management, maximum tolerance in burning is given in the moister sites where severe bums may be required to control competing vegetation. In mesic and drier sites, for the area covered by this diagram, ceanothus seed may be buried in the soil. Therefore, this suggests severe bums should be avoided; spring bums would reduce and delay the gennination and establishment of ceanothus seedlings. In other areas of the ordination, a no-bum recommendation is appropriate to maintain remnant vegetation that can aid in conifer establishment. Such vegetation can act as the target of browsing or may ameliorate the microclimate around planted seedlings. Because response to burning can have a different value for different. objec­ tives, any treatment recommendations should be kept separate by objective. Such a system would enable land managers to weigh the options for each objec­ tive and develop a prescription. This procedure would not only provide a place for managers to record their site· and objective-specific observations and recom· mendations but also show where sound data exist or where speculation is high. The relation between treatment and plant community development is generally vague in the literature as a whole, because sites studied may cover many types, and results are often combined and thus confounded. Integration of existing data into patterns by site type and overall bum severity would prove useful. Acknowledgments W,e gratefully acknowledge those who provided unpublished data for use in this paper: Bill Beese, Louise de Montigny, Michael Feller, Charlie Halpern, and Lance Raff. Useful review comments on this manuscript were provided by Dean DeBell, Charlie Halpern, Donald Hanley, Jerry Kamrnenga, Susan Little, Richard Miller, and Chadwick Oliver. Literature Cited Ahl gren I. F., and C. E. Ahlgren. 1960. Ecological effects of forest fires. Bot. Rev. , 26:483-533. Allen, G. S. 1946. Planning for natural regeneration in the Douglas-fir region. B.C. Lumberman 30(10):53-54, 60. Bigley and Henderson Amo, S. F., D. G. Simmerman, andR. E. Keane. 1985. Forest succession on four habitat types in western Montana. USDA For. Serv. Gen. Tech. Rep. INT-177. Inter­ mountain For. and Range Exp. Stn., Ogden, Utah. 74 p. Austin, R. C., and D. H. Baisinger. 1955. Some effects of burning on forest soils of western Oregon and Washington. J. For. 53(4):275-280. Baker, J. 196 8 . Growth response and chemical composition of Douglas-fiT seedlings on burned and unburned soil samples. Canadian Department of Fisheries and Forestry, For. Res. Lab. Info. Rep. BC- X - 2 8. 20 p. Barnett, D. 1984. Effects off�re on coast range sites. USDA For. Serv. Siuslaw National Forest, WaldportRanger District, Waldpon, Oregon. Bigley, R. E. Forty years of plant succession on slash burned and unburned sites in western Washington and Oregon. In preparation. Bigley, R. E., and R. E. Miller. Effects of slash burning on forest stand establishment and development. In preparation. Boyer, D. E., and J. D. Dell. 1980. Fire effects on Pacific Northwest forest soils. USDA For. Serv. Pacific Northwest Region Watershed Management and Aviation and Fire Management, Portland, Oregon. Braathe, F. 1973. Prescribed burning in Norway: Effects on soil and regeneration. Proceedings, Tall Timbers Fire Ecology Conference 13:211-222. Cattelino, P. J., I. R. Noble, R. 0. Slatyer, and S. R. Kessell. 1979. Predicting the multiple pathways of plant succession. Environ. Manage. 3:41-50. Connell, J. H., and R. 0. Slatyer. 1977. Mechanisms of succession in narural com­ munities and their role in community stability and organization. Am. Nat. 111:1119-1144. de Montigny, L. 1985. The effects of prescribed fire on nill'Ogen dynamics in the Coastal Western Hemlock Zone of B.C. M.F.S. thesis, Yale School of Forestry and Environmental Studies, New Haven, Connecticut. de Montigny, L. E. M., and A. N. D. Auclair. 1982. Fire- productivity interactions: The relation of f1re to the assart effect. In A. N. D. Auclair (ed.) Problem analysis of prescribed burning: A synthesis text. Pac. For. Res. Centre, Victoria, British Columbia. Dymess. C. T. 1965. The effect of Jogging and slash burning on understory vegetation in the H.J. Andrews Experimental Forest. USDA For. Serv.Res. Note PNW 31. Pac. Northwest For. and Ra:_tge Exp. Stn., Portland, Oregon. 13 p. - -- . 1973. Early stages of plant succession following Jogging and burning in the western Cascades of Oregon. Ecology 54(1):57 -69. Effects on Plant Succession Feller, M. C. 1982. The ecological effects of slashbuming with particular reference to British Columbia: A literature review. B.C. Min. For. Land :V1anage. Rep. 13. 60 p. Franklin. J. F. 1982. Forest succession research in the Pacific Northwest: An overview. In J. E. Means (ed.) Forest succession and stand development research in the Northwest, pp. 164-170. Forest Research Laboratory, Oregon State University, Corvallis. Gockerell, E. C. 1966. Plantations on burned versus unburned areas. J. For. 64:392-394. Gratkowski, H. 1961. Brush seedlings after controlled burning of brushlands in south­ west Oregon. J. For. 59:885-888. Grier, C. C. 1972. Effects of fire on the movement and distribution of elements within a forest ecosystem. Ph.D. thesis, University of Washington, Seattle. Halpern, C. B. 1987. Twenty-one years of secondary succession inPseudotsuga forests of the western Cascade Range, Oregon. Ph.D thesis, Oregon State University, Corvallis. . -- Early successional pathways and the resistance and resilience of forest com­ munities. Ecology. In press. --. Early successional patterns of forest species: Interactions of life history traits an d disturbance. Ecology. I n press. Hawkes, B. C. 1986. Micro-plot approach to prescribed ftre effects research. In Proceedings, Prescribed Burning in the Midwest: State of the Art, Symposium held at Stevens Point, Wisconsin. Heavilin, D. 1977. Conifer regeneration on burned and unburned clearcuts on granitic soils of the Klamath National Forest. USDA For. Serv. Res. Note PSW -321. Pac. Southwest For. and Range Exp. Stn., Berkeley, California. 3 p. Hermann, R. K., and W. W. Chilcote. 1965. Effect of seedbeds on germination and survival of Douglas-fir. Oregon State Univ. Res. Pap. 4. 28 p. Ingram, D. C. 193 1. Vegetative changes and grazing use on Douglas-fir cut-over land. I. Agric. Res. 43:387-417. Isaac, L. A. 1940. Vegetative succession following logging in the Douglas-ftr region with special reference to fire. J. For. 38:716-721. -- . 1943. Reproductive habits of Douglas-fir. Charles Lathrop Pack Forestry Foun­ dation. 107 p. Isaac, L. A., and H. G. Hopkins. 1937. The forest soil of the Douglas-fir region, and changes wrought upon it by logging and slash burning. Ecology 18:264-279. Bi£ieY and Henderson Kellman, M. 1974. Preiimina::y seed budgers for two plant communities in coastal British Columbia. Journal of Biogeography 1:123-133. Kienhoiz, R. 1928. Revegetation after logging and bu.""ning in the Douglas-fir region of west= Washington. Illinois State Academy of Science Tr:?..'1Sactions 21:94-108. Knight, H. 1961. Some effecrs of slashburning and clear-cut logging on soil properties and irlltial tree growth. Ph.D. thesis, University of Washington. Seattle. 191 p. Kraemer, J. F. 1977. The long term effecrs of burning on plant succession. M.S. thesis, Oregon State University, Corvallis. Lafferty, R. R. 1980. Post-burn evaluation of plant succession a..."1d forest regeneration in coastal Douglas-fir- western hemlock - red cedar types. Havco Res. Serv. Ltd., Abborsford, British Columbia. Draft report. Lavender, D. P., M. H. Bergman, and L. D. Calvin. 1956. Natural regeneration on staggered settings. Oregon State Board of Forestry Res. Bull. 10. Salem, Oregon. 36 p. Little, S. N., R. D. Ottrnar, and J. L. Ohmann. 1986. Predicting duff consumption from prescribed burns on conifer clearcuts in western Oregon and western Washington. USDA For. Serv. Res. Pap. PNW-362. 29 p. Madison, R. W. 1959. Growth and survival of a Sitka spruce plantation in coastal Oregon. USDA For. Serv. Res. Note 178. Pac. Northwest For. and Range Exp. Stn., Portland, Oregon. 6 p. McCulloch, W. F. 1944. Slash burning. For. Chron. 20:111-118. McLean, A. 1969. Fire resistance of forest species as influenced by root system. J. Range Manage. 22:120-122. Miller. D. L., and D. W. Breuer. 1984. Effects of site preparation by burning and dozer scarification on seedling performance. Potlatch Corporation For. Tech. Pap. TP-84-1. 7 p. Miller, M. 1977. Response of blue huckleberry to prescribed fires in a western Mont.ma larch-fir forest. USDA For. Serv. Res. Pap. IJ'..'T-188. Intermountain For. and Range Exp. Stn., Ogden, Utah. 33 p. Morgan. P., and L. F. Neuenschwander. 1988. Shrub response to high and low seve:-i[)' burns following clearcuuing in northern Idaho. West. J. App. For. 3(1):5-9. Morris, W. G. 1958. Influence of slash burning on regeneration, other plant cover, a."1d flre hazard in the Douglas-fir region. USDA For. Serv. Res. Pap. 29. P2.:.:. 1\onhwest For. and Range Exp. Stn., Portland, Oregon. 49 p. -- . 1970. Effects of slash bu.""ning in overrnature stands of the Douglas-frr region. For. Sci. 16::258-270. 66 Effects on Plant Succession Munger, T. T., and D. N. Matthews. 1941. Slash disposal and forest management after clear cutting in the Douglas-fir region. USDA Circ. 586. Washington, D.C. 55 p. Neal, J. L., E. Wright, and W. B . Bollen. 1 965. Burning Douglas-fir slash: Physical, chemical, and microbial effects in the soil. For. Manage. Res., For. Res. Labora­ tory. Oregon State University, Corvallis. 32 p. Orme, M. L., and T. A. Leege. 1976. Emergence and survival of redstem (Ceanorhus sanguineus) following prescribed burning. In Proceedings, Fourteenth Tall Tim­ bers Fire Ecology Conference and Intermountain Fire Research Council Fire and Land Management Symposium. Missoula, Montana, pp. 391 -420. Tall Timbers Research Station, Tallahassee, Aorida. Pratt, D. W., R. A. Black, and B . A. Zamora. 1984. Buried viable seed in a ponderosa pine community. Can. J. Bot. 62:44-52. Reid, E. H., L. A. Isaac, and G. D. Pickford. 1938. Plant succession on a cut-over, burned, and grazed Douglas fir area. USDA For. Serv. For. Res. Note 26. Pac. Northwest For. and Range Exp. Stn., Portland, Oregon. Roberts, H. A. 1981. Seed banks in soils. Adv. Appl. Bioi. 6 : 1 -55. Ruth. R. H. 1956. Plantation survival and growth in two brush-threat ar eas in coastal Oregon. USDA For. Serv. Res. Pap. 17. Pac. Northwest For. and Range Exp. S tn., Portland, Oregon. 14 p. Shearer, R. C. 1985. Establishment of conifers following broadcast burning on clearcuts in western Montana. Ph.D. thesis, University of Montana, Missoula. 1 15 p. Silen, R. R. 1952. Timing of slash burning with the seed crop: A case history. USDA For. Serv. Res. Note 8 1 . Pac. Northwest For. and Range Exp. Stn., Portland, Oregon. 2 p. Steen, H. K. 1 965. Variarion in vegetation following slash fires near Oakridge, Oregon. USDA For. Serv. Res. Note PNW -25. Pac. Northwest For. and Range Exp. Stn., Portland, Oregon. 6 p. . 1966. Vegetation following slash fires in one western Oregon locality. Northwest Sci. 40: 1 13-120. -- Stein, W. I. 1986. Comparison of site preparation methods on Coast Range sites. USDA For. S erv . Res. Progress Rep. Photocopy. Tarrant, R. E., and E. Wright. 1955. Growth of Douglas-fir seedlings after slashburning. USDA For. Serv. Res. Note PNW- 1 1 5 . Pac. Northwest For. and Range Exp. Stn., Portland, Oregon. 3 p. Topik, C., N. M. Halverson, and D. G. Brockway. 1986. Plant association and manage­ ment guide for the Western Hemlock Zone. USDA For. Serv. R6-ECOL-230A. Pac. Northwest Region, Portland, Oregon. Bio-Jev and Henderson Vihnanek, R. E., and T. M. B allard. 1988. Slashbuming effects on stocking, growth and nutrition of young Douglas-frrplantations in salal-dominated ecosystems of eastern Vancouver Island. Can. J. For. Res. 18:71 8-722. Volland, L. A, and J. D. Dell. 198 1 . Fire effects on Pacific Northwest forest and range vegetation. USDA For. Serv. R6-Rm-067. Pac. Northwest Region, Portland, Oregon. 23 p. Vyse, A. H . and S. J. Muraro. 1973. Reduced planting costs: A prescribed fire benefit. Can. Forestry Serv. Info. Rep. BC-X-84. Pacific Forestry Centre, Victoria, British Columbia. . Weatherspoon, C. P. 1985. Preharvest burning for shrub control in a white fir stand: Preliminary observations. In Proceedings, S ixth Annual Forest Vegetation Management Conference, Redding, California, pp. 7 1 -88. West, N. E., and W. W. Chilcote. 1968. Senecio sylvaticus in relation to Douglas-fir clearcut succession in the Oregon Coast Range. Ecology 49: 1 101-1 107. Yerkes, V. P. 1960. Occurrence of shrubs and herbaceous vegetation after clearcutting old-growth Douglas-frr stands in the Oregon Cascades. USDA For. Serv. Res. Pap. 34. Pac. Northwest For. and Range Exp. Sm., Portland, Oregon. 12 p. 68