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Fire and the Forest History of the North Cascade Range Author(S): Les C

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Fire and the Forest History of the North Cascade Range Author(S): Les C Fire and the Forest History of the North Cascade Range Author(s): Les C. Cwynar Reviewed work(s): Source: Ecology, Vol. 68, No. 4 (Aug., 1987), pp. 791-802 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/1938350 . Accessed: 29/11/2011 12:08 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology. http://www.jstor.org Ecology, 68(4), 1987, pp. 791-802 © 1987 by the Ecological Society of America FIRE AND THE FOREST HISTORY OF THE NORTH CASCADE RANGE1 LES C. CWYNAR2 Departmentof Botany, Universityof Toronto,Toronto, Ontario M5S IA1, Canada Abstract. Postglacial vegetation changes are often ascribed to the direct effects of climate change. I studied pollen, plant macrofossils, and sediment charcoal in order to determine the potential role of changes in the disturbance (fire) regime in the postglacial development of local vegetation at Kirk Lake in the foothills of the North Cascade Range in northwestern Washington. Five pollen assemblage zones are recognized: a Pinus-Populus zone > 12 000 BP, a Picea-Alnus sinuata zone from > 12 000 to 11 030 BP, an Alnus rubra-Pteridium zone from 11 030 to 6830 BP, a Cupressaceae zone beginning at 6830 BP, and a late Holocene Pinus-Alnus rubra zone from 2400 to 900 BP. The first forests (> 12 000 BP) were an open mixture of conifers and deciduous trees, chiefly Tsuga mertensiana, Abies, Pinus contorta, and Populus on a landscape subject to erosion. Just before 12 000 BP, the pioneer species Picea sitchensis, Alnus rubra, and A. sinuata became important constituents of the forest. Although pollen accumulation rates were high, the abundance of Alnus sinuata indicates an open-canopy forest. Beginning 11 200 BP, climatic warming initiated major changes in forest composititon and the fire regime. Tsuga heterophylla migrated into the region, rapidly expanded, then declined shortly thereafter, while Pseudotsuga menziesii, Alnus rubra, and Pteridium expanded, and Pinus contorta, Picea sitchensis, Populus, and Alnus sinuata declined. The abundance of Pseudotsuga, Alnus rubra, and Pteridium between 11 030 and 6830 BP corresponds with increased influxes of charcoal into the sediment; this zone is interpreted as a closed forest with a relatively high fire frequency and composed of a mosaic of postfire successional communities in which fire-adapted Pseudotsuga and Alnus rubra predominated over fire- sensitive Tsuga heterophylla. Pinus monticola became locally important m8000 BP. Be- tween 6800 and 6400 BP Thuja plicata arrived, Tsuga heterophylla expanded, and Alnus rubra, Pseudotsuga, and Pteridium declined. These changes are accompanied by a reduced fire frequency, inferred from lower charcoal accumulation rates, and they indicate a shift to wet-temperate climate similar to today's. The late Holocene fossil record shows the development of the adjacent peatland, which Pinus contorta eventually invaded. Key words: CascadeRange; charcoal;fire; forest history;paleoecology; palynology; plant macro- fossils; Quaternarystudies. INTRODUCTION teraction of physical and biological processes on three time x Recent studies of vegetation emphasize the impor- scales: micro (1-5000 yr, 1-106 m2), macro (5 tance of disturbance (White 1979, Mooney and God- 103-106 yr, 106-1012 m2), and mega (>106 yr, >10'2 ron 1983, Pickett and White 1985). The composition m2). They concluded that "the Quaternary paleoeco- of forests is partly determined by the disturbance re- logical record demonstrates the importance of climatic the gime, which includes average rates, distribution in change as dominant influence upon vegetational space, and severity of disturbances (Loucks 1970, processes and patterns at the macro-scale of spatial- Mooney and Godron 1983, Shugart 1984, Pickett and temporal resolution" and that "even at the micro-scale, White 1985, Runkle 1985). In forests, disturbances prevailing disturbance regimes that affect plant succes- occur on various scales, ranging from small gaps cre- sion must be viewed within the context of their pre- ated by the death of one or a few trees, leading to gap- vailing macroclimate." When macroclimate changes, phase regeneration, to large gaps created by a variety vegetation may therefore be expected to respond not of external factors, such as fire or wind, leading to only to the direct effects of change in climate, but also secondary succession (Runkle 1985). to the indirect effects of change in the disturbance re- Delcourt et al. (1983) developed an hierarchical gime. model of how vegetation patterns arise from the in- Although fire is an important agent of disturbance in the modern evergreen coniferous forests of the Pa- Manuscriptreceived 8 April 1986; revised 15 September cific Northwest and some forest 19 1986. (PNW) types depend 1986; accepted September on it for their and 2 Presentaddress: Department of Biology,Sir Wilfred Gren- regeneration (Franklin Dyrness 1973, fell College, Memorial University of Newfoundland, Corer Agee 1981, Hemstrom and Franklin 1982), the post- Brook, NewfoundlandA2H 6P9, Canada. glacial development of PNW forests is generally inter- 792 LES C. CWYNAR Ecology, Vol. 68, No. 4 preted as resulting from the direct effects of climate the maximum extent of the Cordilleran Ice Sheet (Waitt change. The general sequence of postglacial vegetation and Thorson 1983) of the Fraser Glaciation (circa change at lowland sites in the northern Puget Trough 25 000-10 000 BP). The North Cascades near Glacier (Hansen 1938, 1947, 1948, Hansen and Easterbrook Peak were deglaciated before 12 250 BP (Beget 1984). 1974, Baker 1983, Heusser 1983, Barnosky 1984) be- The vicinity of Kirk Lake was deglaciated before gins with open forests of various admixtures of tem- I 1 500 BP, and the lake probably formed as a kettle perate lowland (Pinus contorta, Populus) and montane in outwash or kame terrace (J. E. Beget, personal taxa (Abies lasiocarpa, Tsuga mertensiana). At - 11 200 communication). Glaciers in the region re-advanced in BP temperate lowland species began to replace mon- the early Holocene 8400-8300 BP (Beget 1984). No tane species so that by 10 000 BP open forests domi- bedrock is exposed within the drainage basin of Kirk. nated by Pseudotsuga and Alnus rubra had developed Surrounding mountains are composed of crystalline at most sites. These forests persisted until -6000 BP rocks, principally schists (McKee 1972). when modern closed forests of temperate conifers The lake lies within the Tsuga heterophylla Zone (Pseudotsuga, Thuja plicata, Tsuga heterophylla) de- (Franklin and Dyrness 1973), which regionally is dom- veloped. The climates inferred from this sequence are inated by Pseudotsuga menziesii (Douglas-fir), Tsuga cool-maritime conditions initially, gradually warming heterophylla (western hemlock), and Thuja plicata to a warmer and drier period relative to modern cli- (western red cedar). Abies amabilis (Pacific silver fir), mate between 10 000 and 6000 BP, and then the de- A. grandis (grand fir), Picea sitchensis (Sitka spruce), velopment of the modern humid climate after 6000 Pinus contorta (lodgepole pine), and P. monticola BP. (western white pine), are minor constituents. The most The role of fire in either maintaining plant com- common deciduous tree is Alnus rubra (red alder). Near munities or in initiating vegetation change has increas- the lake, Thuja plicata and Tsuga heterophylla are the ingly attracted the attention of Quaternary paleoecol- most common trees together with some Pseudotsuga ogists, especially since charcoal is abundant in lake menziesii. A peatland on the east side supports a stand sediments and easily quantified (Waddington 1969, of Pinus contorta with scattered Picea sitchensis and a Swain 1973, Cwynar 1978, Green 1981, Grimm 1983, shrub understory of Ledum groenlandicum (Labrador Lamb 1985, Winkler 1985a, b, Dunwiddie 1986). Some tea) and some Alnus sinuata (Sitka alder). A small stand researchers have recognized the potential significance of Tvpha latifolia (cattail) grows at the north end of of fire in the postglacial development of PNW forests the lake. (Hansen 1947, Heusser 1973a, Mathewes 1973, Leo- pold et al. 1982), but only Sugita and Tsukada (1982) METHODS and Dunwiddie (1986) have provided detailed studies I collected a 12.44-m core from peat on the west of charcoal from lake sediments, and Mathewes (1985) shore of the lake with a 5 cm diameter modified Liv- has stressed the need for direct evidence of past fire ingstone piston sampler (Wright 1967) and help. I re- regimes from sedimentary charcoal profiles for a full moved samples of 0.5 cm3 with a calibrated brass sam- understanding of postglacial vegetation change in the pler (Birks 1976) for pollen analysis (at 20-cm intervals PNW. My primary objective in this paper is to recon- above 9 m, and 5- or 10-cm intervals below 9 m) and struct the history of local vegetation around Kirk Lake, to determine percent mass-loss on ignition (LOI) at Washington in relation to disturbance by fire. I spe- 550°C. Tablets of Eucalyptus pollen or Lycopodium cifically address the following questions: (1) Are post- spores were added to pollen samples for determination glacial changes in vegetation associated with changing of pollen concentration and pollen accumulation rates fire regimes? (2) What was the nature of the vegetation (PARs). I prepared samples for pollen analysis using between 10 000 and 6000 BP and what was the relative standard methods (Faegri and Iversen 1975) and, in importance of change in climate vs. the disturbance addition, sieved basal inorganic sediments to remove regime in initiating the development and maintenance fine particles (Cwynar et al.
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