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Influence of Fire Interval and Serotiny on Postfire Lodgepole Pine Density in Yellowstone National Park

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Influence of Fire Interval and Serotiny on Postfire Lodgepole Pine Density in Yellowstone National Park Utah State University DigitalCommons@USU Quinney Natural Resources Research Library, The Bark Beetles, Fuels, and Fire Bibliography S.J. and Jessie E. 2003 Influence of Fire Interval and Serotiny on Postfire Lodgepole Pine Density in Yellowstone National Park Tania Schoennagel Monica G. Turner William H. Romme Follow this and additional works at: https://digitalcommons.usu.edu/barkbeetles Part of the Ecology and Evolutionary Biology Commons, Entomology Commons, Forest Biology Commons, Forest Management Commons, and the Wood Science and Pulp, Paper Technology Commons Recommended Citation Schoennagel, T., Turner, M. and Romme, W. (2003). Influence of fire interval and serotiny on postfire lodgepole pine density in Yellowstone National Park. Ecology, 84(11): 2967—2978. This Article is brought to you for free and open access by the Quinney Natural Resources Research Library, S.J. and Jessie E. at DigitalCommons@USU. It has been accepted for inclusion in The Bark Beetles, Fuels, and Fire Bibliography by an authorized administrator of DigitalCommons@USU. For more information, please contact [email protected]. Ecology, 84(11), 2003, pp. 2967±2978 q 2003 by the Ecological Society of America THE INFLUENCE OF FIRE INTERVAL AND SEROTINY ON POSTFIRE LODGEPOLE PINE DENSITY IN YELLOWSTONE NATIONAL PARK TANIA SCHOENNAGEL,1,3 MONICA G. TURNER,1 AND WILLIAM H. ROMME2 1Department of Zoology, University of Wisconsin, Madison, Wisconsin 53706 USA 2Department of Forest Sciences, Colorado State University, Fort Collins, Colorado 80523 USA Abstract. The time interval between stand-replacing ®res can in¯uence patterns of initial post®re succession if the abundance of post®re propagules varies with pre®re stand age. We examined the effect of ®re interval on initial post®re lodgepole pine (Pinus contorta var. latifolia Engelm.) density in Yellowstone National Park (YNP) following the 1988 ®res. We asked whether post®re propagule abundance, measured as pre®re percent serotiny, varied with ®re interval and could explain patterns in post®re succession. The response of lodgepole pine density to variation in ®re interval was explained by spatial and temporal variation in pre®re serotiny. At low elevations, post®re lodgepole pine recruitment correlated strongly with pre®re percent serotiny, which varied nonlinearly with pre®re stand age. As a result, post®re lodgepole pine densities varied nonlinearly with ®re interval. In contrast, at high elevations serotiny was low, varied little with stand age and did not in¯uence post®re lodgepole pine densities, although, ®re interval was still a signi®cant predictor of post®re densities. At high elevations, ®re interval varied nonlinearly with post®re lodgepole densities, presumably due to the temporal variation in propagule abundance from open cones in adjacent unburned stands. Temporal variation in stand-level serotiny at low ele- vations was best explained by age of individual trees. Logistic regression indicated that trees expected to be serotinous had a low probability of exhibiting serotiny at a young age, with increasing probability as trees matured up to 140 yr. This increase in serotiny with tree age likely accounts for the initial increase in stand-level percent serotiny with stand age at low elevations. The spatial variation in serotiny was correlated with variation in historical ®re regimes. Fire interval models derived from lower elevations in YNP indicate that ®re occurred historically at 135±185-yr intervals, whereas at higher elevations ®res occurred at 280±310-yr intervals. The spatial patterns of serotiny appear to have been in¯uenced by variability in historical ®re regimes across the Yellowstone landscape, which has conditioned contemporary successional responses to disturbance. Key words: climate change; disturbance; ®re ecology; ®re interval; landscape ecology; multiple successional pathways; Pinus contorta var. latifolia; post®re succession; serotiny; Yellowstone National Park. INTRODUCTION The abundance and arrangement of postdisturbance Fire regimes vary spatially across heterogeneous propagules (viable seeds) in¯uence post®re community landscapes and temporally as climate and fuel condi- composition and structure (Turner et al. 1998). Life tions change. In Yellowstone National Park, mean ®re history traits such as the timing of seed production or intervals (the time between two ®re events at a partic- the time interval required between successful vegeta- ular location) on the subalpine plateaus have varied tive reproduction events, will in¯uence how variation from 70 to 400 yr throughout the Holocene in close in ®re interval affects propagule abundance and post®re association with regional climate variation (Whitlock recruitment. In Yellowstone, the abundance of post®re 1993, Millspaugh et al. 2000). Strong correlations be- propagules is closely tied to serotiny (Tinker et al. tween ®re interval and regional climate have been well 1994, Turner et al. 1997, 1999). Serotiny is the long- documented in many crown-®re dominated systems term storage of seeds within mature closed cones (Cwynar 1987, Clark 1988, 1990, Johnson et al. 1990, which, when heated by ®re, open to release seeds onto Johnson and Wowchuk 1993). Changes in ®re interval newly exposed mineral soil. Lodgepole pine (Pinus may in¯uence successional patterns, yet the direct ef- contorta var. latifolia Engelm.) predominantly exhibit fects of varying ®re intervals on post®re vegetation either open or closed cones (Koch 1996), and we refer have not been well studied in stand-replacing ®re sys- to trees with the ability to produce closed cones during tems. their lifetime as serotinous trees. The proportion of lodgepole pine trees bearing serotinous cones within a Manuscript received 6 May 2002; revised 11 February 2003; stand varies considerably within Yellowstone National accepted 6 March 2003. Corresponding Editor: S. Lavorel. 3 Present address: Department of Geography, University Park (Tinker et al. 1994) and throughout the Rocky of Colorado, Boulder, Colorado 80306 USA. Mountain region (Critch®eld 1957, Muir and Lotan 2967 2968 TANIA SCHOENNAGEL ET AL. Ecology, Vol. 84, No. 11 1985, Koch 1996). Observations also suggest that stand-level proportions of serotiny vary with age of the stand (Mason 1915, Crossley 1956, Lotan 1975). Be- cause viable seeds from multiple years accrue in the canopy of serotinous trees, the density of post®re seed- lings may be linked to the age of stands as they burn (the ®re interval). Pre®re percent serotiny, the percentage of serotinous trees in a stand, has been shown to be a strong predictor of post®re lodgepole pine densities in Yellowstone Na- tional Park (Tinker et al. 1994, Turner et al. 1997, 1999), but how stand-level percent serotiny may vary with time is not well understood. Fire interval relative to levels of serotiny may determine the rate and density of post®re tree recruitment. If ®re occurs in stands when serotiny is low, post®re recruitment may be sparse, as seeds will be consumed rather than released by ®re. If the same site were subject to ®re when levels of ser- otiny are high, post®re recruitment may be very dense. Thus, temporal variation in propagule abundance could be important for predicting successional responses to altered ®re regimes in coniferous forests. If global cli- mate change causes ®re intervals to shift beyond certain FIG. 1. Map of sampling locations and delineation of data temporal thresholds, nonlinear vegetation responses sets analyzed. Triangles represent 25 locations where re- sponse to ®re interval (n 5 50) and pre®re percent serotiny may produce unexpected patterns of succession (Turner at the stand-level were sampled (n 5 50). Circles represent et al. 1998). 10 sites sampled to determine how serotiny varies with tree We studied the effect of ®re interval on initial post- age. Two sites not represented on the map are in high-serotiny ®re succession in Yellowstone following the 1988 ®res areas of Grand Teton National Park. Gray areas represent the high-elevation (dark) and low-elevation (light) regions of the and addressed four questions: (1) Is ®re interval sig- time-since-®re map analyzed to determine historical in¯uence ni®cant in explaining variation in post®re lodgepole of mean ®re intervals on spatial pattern of serotiny. Stippled pine density given similar environmental conditions areas represent lakes. Lines mark all roads and the Yellow- and across broad environmental gradients? (2) How stone National Park boundary. does stand-level serotiny vary with stand age? (3) What mechanisms can explain temporal variation in stand- level serotiny? (4) Does variation in historical ®re re- plateaus (Christensen et al. 1989, Turner et al. 1994). gimes across the landscape explain spatial variation in Fires of this scale have occurred in Yellowstone in the serotiny across the park? past, most recently in the early 1700s (Romme 1982, Romme and Despain 1989). Smaller ®res have also STUDY AREA occurred; for example, within the last century, 4000± Yellowstone encompasses 9000 km2 primarily in the 7000 ha burned yearly in 1910, 1931, 1940, and 1981. northwest corner of Wyoming. Approximately 80% of Although the mean ®re interval for the subalpine pla- the park is covered with coniferous forests dominated teaus is estimated to be ;300±350 yr (Romme 1982), by lodgepole pine (Despain 1990). Our study focused ;32 000 ha of forest on the subalpine plateaus burned on the subalpine forested plateaus that comprise most twice within the last 100 yr, with some stands expe-
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