Surface Temperatures and Durations Associated with Spring Prescribed Fires in Eastern South Dakota Tallgrass Prairies Author(s): Michelle K. Ohrtman, Sharon A. Clay, and Alexander J. Smart Source: The American Midland Naturalist, 173(1):88-98. 2015. Published By: University of Notre Dame DOI: http://dx.doi.org/10.1674/0003-0031-173.1.88 URL: http://www.bioone.org/doi/full/10.1674/0003-0031-173.1.88 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Am. Midl. Nat. (2015) 173:88–98 Surface Temperatures and Durations Associated with Spring Prescribed Fires in Eastern South Dakota Tallgrass Prairies MICHELLE K. OHRTMAN,1 SHARON A. CLAY AND ALEXANDER J. SMART Departments of Plant Science and Natural Resource Management, South Dakota State University, Brookings, 57007 ABSTRACT.—Fire and grazing are commonly used to manage nonnative grasses in the Northern Great Plains, but the effects of fire frequency and management between fire events on fire behavior in this region are poorly understood. We examined temperature and duration of prescribed spring fires at two locations where plots were treated with two fire frequencies (annual or biennial), simulated grazing (1 mo of weekly clipping in spring) and no simulated grazing. In May 2011 and 2013, soil surface fire temperatures and heat duration were monitored in treatment plots using thermocouples. Probes also were placed at 1, 2, and 3 cm depths to measure soil heat transfer. Lethal heat duration (.60 C) at the surface tended to be longer in plots treated with biennial fires compared to plots treated with annual fires. Fires in 2011 had higher maximum temperatures than 2013. Cooler fires in 2013 were characterized by longer durations of lethal heat. However, simulated grazing increased residence time of lethal temperatures in biennial plots and reduced lethal temperature duration in annual plots. Surface heat did not influence soil temperatures even at the 1 cm depth. Greater fuel loads, characteristic of plots treated with biennial fires, generally were associated with higher maximum temperatures and longer heat durations. Results suggest decreasing fire frequency to once every 2 y, perhaps combined with biennial grazing management, may enhance fire behavior to better meet management objectives. INTRODUCTION More than 20,000 ha of grasslands are managed with prescribed fire each year in the Northern Great Plains (NGP), for the primary purpose of exotic plant control and native vegetation enhancement (Willson and Stubbendieck, 1997; NPS, 2009; GPC, 2009, 2010; USFWS, 2010). While grassland fire temperature and duration can, in part, determine the level of plant tissue damage from lethal heat (.60 C) (Whelan, 1995), their effect can be highly variable depending on interactions between numerous abiotic and biotic factors (Pyke et al., 2010). Management (e.g., grazing) between fire events can undoubtedly impact fire behavior in NGP tallgrass prairies, yet few studies have quantified these effects. Fuel quantity and type are among the best predictors of plant tissue heat damage during grassland fires (De Luis et al., 2004; Savadogo et al., 2007; Pyke et al., 2010) with greater fuel loads having consistently higher maximum temperatures and longer plant lethal temperature (.60 C) durations (Stinson and Wright, 1969; Morgan, 1999; Savadogo et al., 2007; Vermeire and Rinella, 2009; Fidelis et al., 2010; Vermeire and Roth, 2011). Fuel moisture content can also impact fire performance (Leonard, 2009). For example dormant vegetation (dry) produced a fire intensity exceeding 700 C for up to 2 min (Stronach and McNaughton, 1989; Stocks et al., 1996) whereas mixed dormant and growing vegetation (higher water content) only reached 100 C for 2 to 3 min durations (Savadogo et al., 2007; Fidelis et al., 2010). Repeated fire events in grasslands are recommended to minimize nonnative plant reinvasion or promote recovery after the initial event (Whisenant, 1990; Kyser and DiTomaso, 2002). However, prescribed grassland fires can remove .99% of aboveground biomass 1 Corresponding author present address: Plant Science Department, South Dakota State University, Brookings, South Dakota 57007. e-mail: michelle.ohrtman@sdstate.edu. 88 2015 OHRTMAN ET AL.: SPRING FIRE BEHAVIOR IN NORTHERN MANAGED PRAIRIE 89 (Bragg, 1982; Howe, 1994b) and annual fire frequencies may not have sufficient fuel loads present to provide the necessary dose of lethal heat for suppression (NPS, 2009). Many studies suggest grassland fuel loads do not accumulate in a linear manner. Vegetation biomass can reach prefire levels in a minimum of 2 y (Morgan, 1999; Prober et al., 2007; Fidelis et al., 2010; Kirkpatrick et al., 2011; MacDonald and McPherson, 2011), although little or no further build up may occur if return times are greater than four or more years (Govender et al., 2006; Oluwole et al., 2008; Cianciaruso et al., 2010). Grazing management between fire events is another important variable that can impact the success of a fire treatment. Grazing used in combination with fire increases grassland species diversity (Collins et al., 1998) and effectively manages invasive plants (Davies et al., 2009; Diamond et al., 2009; MacDonald and McPherson, 2011). Although grazing removes less plant biomass than fire, grazed lands have lower fuel loads than ungrazed areas (Savadogo et al., 2007; Davies et al., 2009; Diamond et al., 2009; Leonard et al., 2010; Kirkpatrick et al., 2011; MacDonald and McPherson, 2011), which may reduce the initial effectiveness of fire to kill undesirable vegetation. The combination of frequent fires, with grazing between fire events, can further reduce post treatment fuel accumulation (Archibald et al., 2005; Kirkpatrick et al., 2011); the unintended consequence can be milder fire conditions compared to areas managed with either method alone. Similar to fire, the effects of grazing management on fuel loads may be negligible beyond 2 to 3 y (Archibald et al., 2005; Kirkpatrick et al., 2011; MacDonald and McPherson, 2011). Spring prescribed fires are well known to reduce exotic cool season grass cover when set during their elongation phase of growth by damaging plant meristems through prolonged exposure to lethal heat (Willson and Stubbendieck, 1997). Consequently, spring fires can promote dominance of native warm season grasses by altering competitive interactions between plant species (Howe, 1994a, b, 1995; Smith and Knapp, 1999; Howe, 2000). In addition spring fires remove thatch and standing dead biomass which can enhance warm season grass growth by allowing greater levels of light at the soil surface and higher soil temperatures early in the growing season (Knapp and Seastedt, 1986). Both annual (Abrams and Hulbert, 1987; Abrams, 1988; Towne and Kemp, 2003) and biennial (Svedarsky et al., 1986; Towne and Kemp, 2008) spring fires have achieved these management goals. In the North American Great Plains, a region where the fire and grazing interaction is considered a single ecological process (Hamilton, 2007; Fuhlendorf et al., 2008), no research has examined the combined effects of fire frequency and grazing on fire behavior. We investigated the influence of spring fire return frequency and simulated spring grazing treatments on fire temperature and lethal heat duration at two locations in eastern South Dakota. We hypothesize: (1) biennial fires will produce higher maximum temperatures and longer durations of plant lethal temperatures than annual fires and (2) post fire simulated grazing will decrease maximum fire temperature and lethal heat duration. Research findings can be used to better understand how grazing and prescribed fires in northern tallgrass prairies influence fuel dynamics and fire behavior to optimize management effects. METHODS STUDY SITES This research was performed at two sites in Brookings County, located in eastern South Dakota. The first site, nearest to the city of Brookings (44u2096.330N, 96u48928.620W), was a cool season pasture that had been seeded with big bluestem (Andropogon gerardii Vitman) in 2005. Soil at the Brookings site was well drained Barnes clay loam with 0 to 2% slopes (NRCS, 2010a, b). Kentucky bluegrass (Poa pratensis L.), big bluestem, and smooth 90 THE AMERICAN MIDLAND NATURALIST 173(1) bromegrass (Bromus inermis Leyss. subsp. inermis) were the dominant grass species and yellow sweetclover [Melilotus officinalis (L.) Lam.] was the predominant forb species. Average vegetation biomass in untreated areas at this site in 2011 was 3400 kg ha21. The second site, near the city of Volga (44u2391.530N, 96u57929.390W) was a native prairie with well drained soils in the Buse Poinsett complex with 9 to 15% slopes (NRCS, 2010a, b). Common plant species at the Volga site included little bluestem [Schizachyrium scoparium (Michx.)], big bluestem, sideoats grama [Bouteloua curtipendula (Michx.) Torr.], smooth bromegrass, and yellow sweetclover. Kentucky bluegrass was also present but less prevalent than at the Brookings site. Average vegetation biomass collected in untreated plots at this site in 2011 was 2300 kg ha21. Annual precipitation recorded at Brookings, South Dakota was 41.5 and 46.8 cm in 2011 and 2013, respectively (South Dakota Climate and Weather, 2013). Field plots were established in 2009 in a split block – split plot design containing four replicates of each treatment at each site (Brookings and Volga). Fire treatments were applied to 6 3 6 m blocks and consisted of no fire (control), annual spring fire, and biennial spring fire. Each block was split into two 3 3 6 m plots and one plot of each block was treated with simulated grazing. Simulated grazing consisted of clipping vegetation to 2 cm height once each week for 4 wk following the fire treatments; the first clipping treatment occurred 7 d after fire. Clipped biomass was removed from the study plots. All prescribed fires were conducted in May using a ring fire method when there was a mix of dormant and actively growing vegetation that is common during spring prescribed fires in the NGP. Weather conditions during fire treatments consisted of wind speeds between 8 and 29 kph, air temperatures between 10 and 24 C, and relative humidities between 30 and 50%. Biennial prescribed fires and the subsequent simulated grazing treatments were applied in 2009, 2011, and 2013. Annual prescribed fires and postfire simulated grazing treatments were applied in 2009, 2010, 2011, 2012, and 2013. FUEL CONDITIONS In 2011 fuel load was quantified in all treatment plots (biennial and annual fire with and without simulated grazing) at the Brookings site and in annual fire treatments at Volga. Live and dead herbaceous vegetation was clipped to ground level within 0.25 m2 quadrats (n 5 4 treatment21) and dried for 3 d at 70 C. Fuel weights were recorded immediate following collection and after drying to determine percent moisture. In 2013 fuel load was quantified only in plots without simulated grazing from both the Brookings and Volga sites; however fuel moisture was not measured. Surface soil (0 to 7.5 cm depth) moisture values were obtained within treatment plots but found to be negligible, and therefore these data are not discussed. MAXIMUM TEMPERATURE AND LETHAL HEAT DURATION Surface and subsurface temperature was measured during prescribed fires on May 6, 2011 and May 16, 2013 at both study sites. To measure surface temperatures during prescribed fires, mineral insulated thermocouples (Type K) attached to data loggers (TC Direct, Hillside, IL) were placed within the surface litter layer taking care not to embed probes in the soil. However, litter depth varied by plot and site and therefore probes recorded within a range of 5 cm above the soil surface. Four probes were placed in each plot within three of the four blocks (12 probes total for three replicates) for each site. However, probes were often disturbed during fire ignition and containment, leaving two or three probes per replicate (seven to nine probes total) with accurate temperature recordings treatment21 site21 year21. Thermocouples were placed at least 1 m inside the plot boundary and spaced at least 1 m apart. Temperatures were recorded at 1 s intervals from just prior to fire ignition 2015 OHRTMAN ET AL.: SPRING FIRE BEHAVIOR IN NORTHERN MANAGED PRAIRIE 91 until surface temperatures were below 60 C. Subsurface temperatures were monitored in one block at each site by installing thermocouples at 1, 2, and 3 cm depth (n 5 3 thermocouples depth21) in the vertical face of a shallow pit. However, no significant change in subsurface soil temperature was observed during annual and biennial fires so results are not shown. STATISTICAL ANALYSIS Mixed effects ANOVA models and student’s t-tests were used to test the effects fire return frequency (annual or biennial), simulated grazing treatment (postfire clipping or no clipping), location (Brookings and Volga), and year (2011 and 2013) on maximum fire temperature and duration of plant lethal heat (.60 C). Site, fire frequency, and simulated grazing were considered fixed effects and year and block were considered random effects. Differences among treatments were also examined for fuel load and fuel moisture by location and year. Means (6SE) are reported with significant differences separated using Fisher’s Protected LSD Test when P ,0.05. Analyses were performed in JMP 10.0.2 (SAS Institute Inc.). RESULTS FUEL CONDITIONS Brookings.—Fuel loads in 2011 were 3700 6 670 kg ha21 in plots treated with biennial prescribed fires and 2500 6 310 kg ha21 in plots treated with annual prescribed fires (P 5 0.05). Fuel loads did not differ between clipped and unclipped plots at the time of burning during this year (P 5 0.25), regardless of fire return frequency. Vegetation moisture was about 16% irrespective of fire frequency (P 5 0.36) and simulated grazing treatment (P 5 0.68). In 2013 fuel load was only measured in plots that were not subjected to the simulated grazing treatment. During that year fuel loads were 8000 6 430 kg ha21 in plots treated with biennial prescribed fires and 5200 6 780 kg ha21 in the plots treated with annual prescribed fires. Fuel loads in plots treated with biennial prescribed fires were 35% more than that fuel loads in plots treated with annual prescribed fires, irrespective of year (P 5 0.001). Volga.—There was a trend of lower fuel load in plots treated with simulated grazing than unclipped plots in annual fire treatments at this site in 2011 (2800 6 150 kg ha21 vs. 3500 6 360 kg ha21; P 5 0.07). In 2013 fuel loads in biennial prescribed fire plots at the Volga site were 48% greater than annual plots (4800 6 820 kg ha21 vs. 2500 6 380 kg ha21; P 5 0.004); these values are 35 and 52% less than respective fuel loads at Brookings (P 5 0.002). MAXIMUM TEMPERATURE AND LETHAL HEAT DURATION Maximum fire temperature.—Maximum fire temperature differed by year (P 5 0.001) and there was an interaction between fire return frequency and simulated grazing (P 5 0.05) (Table 1). Averaged across all treatment combinations and sites, maximum fire temperatures in 2011 were approximately 230 C hotter compared to fire temperatures in 2013. Maximum fire temperatures in plots treated with biennial fires but not treated with simulated grazing were higher than in plots treated annual fires and simulated grazing. The highest recorded temperature during our study, 836 C, was recorded during 2011 in a plot treated with biennial fire but not treated with simulated grazing. Lethal heat duration.—Three-way interactions were noted when examining the length of time the surface remained .60 C (Table 1). Lethal heat durations of temperatures .60 C at the Volga site tended to be shorter than at the Brookings site (Fig. 1, Table 2). Higher maximum temperatures were associated with shorter periods of lethal heat duration in plots 92 THE AMERICAN MIDLAND NATURALIST 173(1) TABLE 1.—ANOVA for maximum temperature (R2 5 0.46, F ratio 5 12.79, P , 0.0001) and temperature duration above 60 C (R2 5 0.54, F ratio 5 12.43, P , 0.0001). Degrees of freedom 5 1 for each parameter Maximum temperature .60 C duration Parameter Sum of squares F ratio Sum of squares F ratio year site fire frequency clip year*site year*fire frequency year*clip site*fire frequency site*clip fire frequency *clip year*fire frequency *clip site*fire frequency *clip 1788196.4 19303.5 228701.6 74538.0 105586.0 5739.0 48376.9 27094.2 32294.1 114563.5 - 98.2** 1.1 12.6 4.1 5.8 0.3 2.7 1.5 1.8 6.3* - 348126.1 298528.5 339981.8 2819.5 11178.3 1376.6 5135.1 28108.2 22344.5 399.7 35392.0 29314.6 49.2** 42.2** 48.0** 0.4 1.6 0.2 0.7 4.0 3.2 0.1 5.0* 4.1* Significance: ** ,0.0001, * ,0.05 treated with annual fires (R2 5 0.10, F ratio 5 8.2, P 5 0.006). However, plots treated with both biennial fires and simulated grazing at the Brookings site were characterized by longer periods of lethal heat; this result was not found at the Volga site. At both sites and during both years of measurement, plots treated with annual fires but not treated with simulated grazing had shorter periods of lethal heat compared to plots treated with biennial fires and not treated with simulated grazing. In 2013 at both sites, plots treated with annual fires and simulated grazing had shorter duration of lethal heat compared to plots treated with annual fires but not treated with simulated grazing. In 2013 simulated grazing had inconsistent effects on lethal heat duration when applied to biennially burned plots. Periods of lethal heat duration were longer in 2013 compared to 2011. DISCUSSION Our results support previous reports that document the importance of fuel load for predicting prescribed fire temperature and duration (Stinson and Wright, 1969; Morgan, 1999; Fidelis et al., 2010). Biennial frequency fire plots at Brookings had 30% greater fuels and greater durations above 60 C, that ranged between 60 to 180 s longer than annual plots, irrespective of year. In addition the Brookings site had nearly twice the fuel loads compared with the Volga site and Brookings fires had greater durations of plant lethal temperature (up to 120 s longer) than Volga in both years. The difference in fuel loads between sites can be explained, in part, by an increase in native warm season grass cover at Brookings over time. Big bluestem cover increased from 34 to 80% in annually treated plots and from 19 to 56% in biennial plots since 2009 (Smart et al., 2013). Volga also experienced an increase in native warm season grass cover over time with fire and simulated grazing management but little bluestem and sideoats grama were dominant. Another South Dakota study observed vegetation biomass to be two times greater following prescribed fires where big bluestem was the dominant plant species (.60%) than where this species was absent (Engle and Bultsma, 1984). Frequent prescribed fires have been reported to produce high amounts of native warm season grasses but this management technique can also reduce species diversity 2015 OHRTMAN ET AL.: SPRING FIRE BEHAVIOR IN NORTHERN MANAGED PRAIRIE 93 FIG. 1.—Temperature profiles (above 60 C) for spring prescribed fires at Brookings and Volga, South Dakota in (A) 2011 and (B) 2013 for plots treated with a combination of annual or biennial (every two years) fire and annual simulated grazing (Clip) or no grazing treatment (No Clip). Lines represent an average of 7 to 9 temperature readings per treatment. Note: these are not true averages of high temperature or duration but represent averages from the initial second each probe exceeded 60 C until each probe fell below this temperature (Collins et al., 1998; Kindscher and Tieszen, 1998; Polley et al., 2005) which may not be a desired management response. Flame residence times in North American grasslands have been reported to be about 15 s long with maximum flame temperatures between 800 and 1000 C (Wotton, 2009). Average maximum temperatures at the soil surface in our plots were cooler (Table 2) but durations of plant lethal heat recorded in biennial plots in this study were greater than previous reports. For example annual prescribed grassland fires in Australia were observed to reside above 100 C for 62 s on average, whereas a single replication of fire every 2, 4, and 7 y resided for 68, 113, and 173 s, respectively (Morgan, 1999). Annual fires recorded by this study in 2011 produced comparable heat durations to those reported by Morgan (1999), but temperatures .100 C in annual and biennial plots in 2013 resided between 2 and 4.5 times longer depending on site and simulated grazing treatment. Sites with frequent fire recorded 94 THE AMERICAN MIDLAND NATURALIST 173(1) TABLE 2.—Maximum fire temperature and duration of fire above 60 C for annual and biennial plots by clipping treatment burned in 2011 and 2013 at Brookings and Volga, South Dakota. Values represent an average of three replicates (with two or three temperature readings per replicate) per site. Significant differences are separated using Fisher’s Protected LSD Test when P , 0.05. Means followed by the same letter within a year are not significantly different across sites or across treatments within a site 2011 Annual fire Site Clip No clip Maximum temperature (C) Brookings 438 bc 491 abc Volga 378 c 549 ab Temperature duration (s) .60 C Brookings 177 bc 138 cd Volga 131 cd 83 d Temperature duration (s) .100 C Brookings 91 bc 93 bc Volga 58 c 53 c Temperature duration (s) .200 C Brookings 47 bc 50 bc Volga 29 c 29 c 2013 Biennial fire Annual fire Clip No clip Biennial fire Clip No clip Clip No clip 467 abc 528 abc 523 abc 611 a 205 cd 152 d 336 ab 209 cd 414 a 283 bc 262 bc 290 bc 292 a 151 bcd 286 a 229 ab 255 cde 181 e 305 bc 223 de 442 a 233 cde 371 ab 293 cd 200 a 90 bc 213 a 121 b 142 cde 75 e 191 bc 121 de 306 a 157 cd 242 ab 180 bcd 121 a 56 bc 131 a 76 b 38 bcd 1d 95 b 20 cd 166 a 52 bcd 91 b 68 bc 15 to 235 s of lethal heat during prescribed grassland fires, whereas a site unburned for 6 y produced these temperatures for 140 to 330 s (Fidelis et al., 2010). Stinson and Wright (1969) observed similar durations of temperatures .60 C for head fires in southern mixed prairie of unreported fire frequency. Whereas most of our fires recorded temperature durations within this range, we observed plant lethal heat durations between 250 and 440 s on average during prescribed fires at Brookings in 2013. These data suggest plant lethal temperature duration may be underestimated in grassland fire research if the temperature profile is not monitored. Both biennial and annual fire return frequencies have been successful for reducing cover of cool season exotic grasses (Smart et al., in press) but waiting 2 y between fires may enhance control efforts while reducing costs and providing greater habitat for wildlife (Herkert, 1994; Dechant et al., 2003). Biennial simulated spring grazing combined with biennial fire produced some of the longest durations of plant lethal heat in both years and the highest maximum temperatures in 2013. These results are unexpected because simulated spring grazing treatments removed surface vegetation, leading to lower (but not statistically significant) fuel loads before the fires in clipped than unclipped plots. However, in mesic areas or during wetter years, the interaction between fire and post fire grazing can enhance plant diversity and production (Fuhlendorf and Engle, 2001) and perhaps change the quality of vegetation fuels. Grazing by livestock may have different effects on fire performance than the uniform clipping treatments used in this study. Even at heavy stocking rates, utilization has been reported to be only 60% (Smart et al., 2010) whereas simulated grazing removed all vegetation fuels above 2 cm height. In addition grazing animals selectively remove palatable herbaceous vegetation with higher moisture content and leave dry litter behind; this process can lower the moisture content of the fuel load and increase fire severity (Leonard et al., 2010). However, most combined fire and grazing programs graze within a few weeks to months following the fire when there is less unpalatable dry material to be avoided (Anderson et al., 1970; Kirkpatrick et al., 2011). 2015 OHRTMAN ET AL.: SPRING FIRE BEHAVIOR IN NORTHERN MANAGED PRAIRIE 95 Simulated grazing was only found to increase fire temperature and duration at the Brookings site. Simulated grazing likely provided further setback of cool season species attempting to recover from fire, which would enhance native warm season grass growth through reduced competition. The effects of this combined management may have been more pronounced at Brookings because of the high level of pre management cool season cover and the dominance of more productive big bluestem (Engle and Bultsma, 1984; Smart et al., in press). Post fire grazing in northern grasslands therefore has the potential to increase the ability of future fires to achieve management goals by stimulating vegetation growth immediately after the disturbance and promoting greater accumulation of surface fuels. Fire and grazing are successful for grassland management because native species that evolved with repeated disturbance recover quickly. Plant biomass reduction following fire is often short lived; 2 y after fire and grazing vegetation fuels on all plots were not significantly different (MacDonald and McPherson, 2011). On the other hand, fuel loads can increase with longer periods between fires with greater fuel accumulations 3 y after the fire than pre fire (Ducherer et al., 2009). Indeed, greater time between fires has been observed to translate to greater durations of plant lethal heat (Morgan, 1999). The observed negative correlation between maximum fire temperature and duration of lethal heat for annual fire frequencies supports lighter fuel loads, associated with more frequent fires, burn quickly and maintain shorter durations of plant lethal heat at the soil surface. We did not test the effects of longer periods between fire or grazing ($3 y) on fire behavior in NGP grasslands. However, if longer periods between fire and grazing are used there may be an increased risk of nonnative plant re- invasion or recovery. We also used a ring fire method where backfires burned most of the plot areas in this study; such fires promote lower fire temperatures and durations than headfires (Laterra et al., 2006; Savadogo et al., 2007). With comparable fuel loads, headfires produce higher temperatures and durations that can be more effective for invasive plant control (MacDonald et al., 2007; Vermeire and Rinella, 2009). Different fire regimes (i.e., frequency, timing) may favor different species (Whelan, 1995; Pivello and Norton, 1996; Cianciaruso et al., 2010) and therefore it is important to understand the fire tolerance of target vegetation as well as the capacity of the fire to produce the necessary duration of heat. Although cover responses of Kentucky bluegrass, smooth brome, and native warm season grasses to fire management are well documented, the specific heat tolerance of common tallgrass prairie species at various life stages (from seed to mature plants) is unknown. Reduced cover of Kentucky bluegrass and smooth brome and greater dominance of big and little bluestem suggest repeated late spring prescribed fires at Brookings and Volga provided sufficient energy to suppress exotic cool season grass growth (Smart et al., 2013). However, this study showed prescribed fires did not provide sufficient heat to result in plant lethal heat transfer in the soil. We did not observe an increase in temperature at 1 cm soil depth, irrespective of fire frequency treatment. These results suggest seeds with any soil insulation and established plants with belowground perennating buds will receive minimal damage from prescribed spring grassland fires on lands with recent fire and grazing management (#2 y). Other control methods, in addition to fire, may need to be considered when developing management strategies for unwanted perennial species. This study is the first to describe fire behavior in managed North American tallgrass prairie and suggests fuel load is an important variable for predicting fire behavior in this system. We found biennial fire regimes can extend the duration of plant lethal heat by promoting greater fuel accumulations and therefore may have the greatest potential for 96 THE AMERICAN MIDLAND NATURALIST 173(1) delivering the necessary lethal heat dose for invasive plant control. Combined biennial fire and post fire grazing can also stimulate fuel production but the effects of this management are dependent on site-specific and climatic factors. This study showed fire behavior is variable across locations and years and therefore site specific evaluations over time are important. Acknowledgments.—This research was supported by a grant from USDA/CSREES Rangeland (award 00486094). Additional funding was provided by South Dakota Agricultural Experiment Station and South Dakota Cooperative Extension Service. Thanks to Stephanie Hansen, Jiyul Chang, Shauna Waughtel-Johnson, Megan Mortallero, Emily Helms, and Heidi Myer for field assistance. LITERATURE CITED ABRAMS, M. D. 1988. Effects of burning regime on buried seed banks and canopy coverage in a Kansas tallgrassprairie. Southwest Nat., 33:65–70. ——— AND L. C. HULBERT. 1987. Effect of topographic position and fire on species composition in tallgrass prairie in northeast Kansas. Am. Midl. Nat., 117:442–445. ANDERSON, K. L., E. F. SMITH, AND C. E. OWENSBY. 1970. Burning Bluestem Range. J. Range Manage., 23:81–92. ARCHIBALD, S., W. J. BOND, W. D. STOCK, AND D. H. K. FAIRBANKS. 2005. Shaping the landscape: fire-grazer interactions in an African savanna. Ecol. Appl., 15:96–109. BRAGG, T. B. 1982. Seasonal variations in fuel and fuel consumption by fires in a bluestem prairie. Ecology, 63:7–11. CIANCIARUSO, M. V., I. A. DA SILVA, AND M. A. BATALHA. 2010. Aboveground biomass of functional groups in the ground layer of savannas under different fire frequencies. Aust. J. Bot., 58:169–174. COLLINS, S. L., A. K. KNAPP, J. M. BRIGGS, J. M. BLAIR, AND E. L. SEINAUER. 1998. Modulation of diversity by grazing and mowing in native tallgrass prairie. Science, 280:745–747. DAVIES, K. W., T. J. SVEJCAR, AND J. D. BATES. 2009. Interaction of historical and nonhistorical disturbances maintains native plant communities. Ecol. Appl., 19:1536–1545. DE LUIS, M., M. J. BAEZA, J. RAVENTOS, AND J. GONZALEZ-HIDALGO. 2004. Fuel characteristics and fire behaviour in mature Mediterranean gorse stands. Int. J. Wildland Fire, 13:79–87. DECHANT, J. A., M. L. SONDREAL, D. H. JOHNSON, L. D. IGL, C. M. GOLDADE, M. P. NENNEMAN, AND B. R. EULISS. 2003. Effects of management practices on grassland birds: Grasshopper Sparrow. Northern Prairie Wildlife Research Center, Jamestown, ND. Jamestown, ND. 28 p. http://www.npwrc. usgs.gov/resource/literatr/grasbird/grsp/grsp.htm (Version 12AUG2004). Accessed Feb. 17, 2012. DIAMOND, J. M., C. A. CALL, AND N. DEVOE. 2009. Effects of targeted cattle grazing on fire behavior of cheatgrass-dominated rangeland in the northern Great Basin, USA. Int. J. Wildland Fire, 18:944–950. DUCHERER, K., Y. BAI, D. THOMPSON, AND K. BROERSMA. 2009. Dynamic responses of a British Columbian forest-grassland interface to prescribed burning. West N. Am. Naturalist, 69:75–87. ENGLE, D. M. AND P. M. BULTSMA. 1984. Burning of northern mixed prairie during drought. J. Range Manage., 37:398–401. FIDELIS, A., M. D. DELGADO-CARTAY, C. C. BLANCO, S. C. MULLER, V. D. PILLAR, AND J. PFADENHAUER. 2010. Fire intensity and severity in Brazilian campos grasslands. Interciencia, 35:739–745. FUHLENDORF, S. D. AND D. M. ENGLE. 2001. Restoring heterogeneity on rangelands: ecosystem management based on evolutionary grazing patterns. Bioscience, 51:625–632. ———, ———, J. KERBY, AND R. HAMILTON. 2008. Pyric herbivory: rewilding landscapes through the recoupling of fire and grazing. Conserv Biol., 23:588–598. [GPC] NORTHERN GREAT PLAINS INTERAGENCY DISPATCH CENTER. 2009. 2009 Annual Report. http://gacc.nifc. gov/rmcc/dispatch_centers/r2gpc/2009_Annual_rpt.pdf Accessed Feb. 2, 2012. ———. 2010. 2010 Annual Report. http://gacc.nifc.gov/rmcc/dispatch_centers/r2gpc/2010_Annual_ rpt.pdf Accessed Feb. 2, 2012. 2015 OHRTMAN ET AL.: SPRING FIRE BEHAVIOR IN NORTHERN MANAGED PRAIRIE 97 GOVENDER, N., W. S. W. TROLLOPE, AND B. W. VAN WILGEN. 2006. The effect of fire season, fire frequency, rainfall and management on fire intensity in savanna vegetation in South Africa. J. Appl. Ecol., 23:748–758. HAMILTON, R. G. 2007. Restoring heterogeneity on the tallgrass prairie preserve: applying the fire grazing interaction model [abstract], p. 163–169. In: R. E. Masters and K. E. M. Galley (eds.). Proceedings of the 23rd Tall Timbers Fire Ecology Conference: Fire in grassland and shrubland ecosystems. Bartlesville, OK. Tall Timbers Research, Inc, Tallahassee, FL. HERKERT, J. R. 1994. The effects of habitat fragmentation on Midwestern grassland bird communities. Ecol. Appl., 4:461–471. HOWE, H. F. 1994a. Managing species diversity in tallgrass prairie: assumptions and implications. Conserv. Biol., 8:691–704. ———. 1994b. Response of early and late flowering plants to fire season in experimental prairies. Ecol. Appl., 4:121–133. ———. 1995. Succession and fire season in experimental prairie plantings. Ecology, 76:1917–1925. ———. 2000. Grass response to seasonal burns in experimental plantings. J. Range Manage., 53:437–441. KINDSCHER, K. AND L. L. TIESZEN. 1998. Floristic and soil organic matter changes after five and thirty-five years of native tallgrass prairie restoration. Restor. Ecol., 6:181–196. KIRKPATRICK, J. B., J. B. MARSDEN-SMEDLEY, AND S. W. J. LEONARD. 2011. Influence of grazing and vegetation type on post fire flammability. J. Appl. Ecol., 48:642–649. KNAPP, A. K. AND T. R. SEASTEDT. 1986. Detritus accumulation limits productivity of tallgrass prairie. Bioscience, 36:662–668. KYSER, G. B. AND J. M. DITOMASO. 2002. Instability in a grassland community after the control of yellow starthistle (Centaurea solstitialis) with prescribed burning. Weed Sci., 50:648–657. LATERRA, P., E. Z. ORTEGA, M. D. C. OCHOA, O. R. VIGNOLIO, AND O. N. FERNANDEZ. 2006. Interactive influences of fire and vertical distribution of seed banks on post fire recolonization of a tall tussock grassland in Argentina. Austral. Ecol., 31:608–622. LEONARD, S. W. J. 2009. Predicting sustained fire spread in Tasmanian native grasslands. Environ. Manage., 44:430–440. LEONARD, S. J. KIRKPATRICK, AND J. MARSDEN-SMEDLEY. 2010. Variation in the effects of vertebrate grazing on fire potential between grassland structural types. J. Appl. Ecol., 47:876–883. MACDONALD, N. W., B. T. SCULL, AND S. R. ABELLA. 2007. Mid spring burning reduces spotted knapweed and increases native grasses during a Michigan experimental grassland establishment. Restor. Ecol., 15:118–128. MACDONALD, C. J. AND G. R. MCPHERSON. 2011. Absence of a grass/fire cycle in a semiarid grassland: response to prescribed fire and grazing. Rangeland Ecol. Manage, 64:384–393. MORGAN, J. W. 1999. Defining grassland fire events and the response of perennial plants to annual fire in temperate grasslands of southeastern Australia. Plant Ecol., 144:127–144. [NPS] NATIONAL PARK SERVICE. 2009. Northern Great Plains Fire Management. http://www.nps.gov/ ngpfire/ Accessed Feb. 2, 2012. [NRCS] NATURAL RESOURCES CONSERVATION SERVICE, U.S. DEPARTMENT OF AGRICULTURE. 2010a. Soil Series Classification. https://soilseries.sc.egov. usda.gov/scname.asp. Accessed: Dec. 16, 2010. ———. 2010b. Web Soil Survey. http://websoilsurvey.nrcs.usda.gov/app/WebSoilSurvey.aspx. Accessed: Dec. 16, 2010. OLUWOLE, F. A., S. J. MACKSON, AND S. DUBE. 2008. Long term effects of different burning frequencies on the dry savannah grassland in South Africa. Afr. J. Ag. Res., 3:147–153. PIVELLO, V. R. AND G. A. NORTON. 1996. Fire tool, an expert system for the use of prescribed fires in Brazilian savannas. J. Appl. Ecol., 33:348–356. POLLEY, H. W., J. D. DERNER, AND B. J. WILSEY. 2005. Patterns of plant species diversity in remnant and restored tallgrass prairies. Restor. Ecol., 13:480–487. PROBER, S. M., K. R. THIELE, AND I. D. LUNT. 2007. Fire frequency regulates tussock grass composition, structure and resilience in endangered temperate woodlands. Austral. Ecol., 32:808–824. PYKE, D. A., M. L. BROOKS, AND C. D. ANTONIO. 2010. Fire as a restoration tool: A decision framework for predicting the control or enhancement of plants using fire. Restor. Ecol., 18:274–284. 98 THE AMERICAN MIDLAND NATURALIST 173(1) SAVADOGO, P., D. ZIDA, L. SAWADOGO, D. TIVEAU, M. TIGABU, AND C. ODÉN. 2007. Fuel and fire characteristics in savanna woodland of West Africa in relation to grazing and dominant grasstype. Int. J. Wildland Fire, 16:531–539. SMART, A. J., J. D. DERNER, J. R. HENDRICKSON, R. L. GILLEN, B. H. DUNN, E. M. MOUSEL, P. S. JOHNSON, R. N. GATES, K. K. SEDIVEC, K. R. HARMONEY, J. D. VOLESKY, AND K. C. OLSON. 2010. Effects of grazing pressure on efficiency of grazing on North American Great Plains rangelands. Rangeland Ecol. Manage., 63:397–406. ———, T. K. SCOTT, S. A. CLAY, D. E. CLAY, AND M. OHRTMAN. 2013. Spring clipping, fire, and simulated increased atmospheric nitrogen deposition effects on tallgrass prairie vegetation. Rangeland Ecol. Manage, 66:680–687. SMITH, M. D. AND A. K. KNAPP. 1999. Exotic plant species in a C4 dominated grassland: invasibility, disturbance, and community structure. Oecologia, 120:605–612. STINSON, K. J. AND H. A. WRIGHT. 1969. Temperatures of headfires in the southern mixed prairie of Texas. J. Range Manage., 22:169–174. STOCKS, B. J., B. W. VAN WILGEN, W. S. W. TROLLOPE, D. J. MCRAE, J. A. MASON, F. WEIRICH, AND A. L. F. POTGEITER. 1996. Fuels and fire behavior dynamics on large scale savanna in Kruger National Park, South Africa. J. Geophys. Res., 101:23,541–23, 550. STRONACH, N. R. H. AND S. T. MCNAUGHTON. 1989. Grassland fire dynamics in the Serengeti ecosystem, and a potential method of retrospectively estimating fire energy. J. Appl. Ecol., 26:1025–1033. SVEDARSKY, W. D., P. E. BUCKLEY, AND T. A. FEIRO. 1986. The effect of 13 years of annual burning on an aspen prairie ecotone in northwestern Minnesota, p. 118–122. In: G. K. Clambey and R. H. Pemble (eds.), Proceedings, 9th North American Prairie Conference-The Prairie: Past, Present and Future. Moorhead, MN, and Fargo, ND: Tri College University. TOWNE, E. G. AND K. E. KEMP. 2003. Vegetation dynamics from annually burning tallgrass prairie in different seasons. J. Range Manage., 56:185–192. ——— AND ———. 2008. Long term response patterns of tallgrass prairie to frequent summer burning. Rangeland Ecol. Manage., 61:509–520. [USFWS] UNITED STATES FISH AND WILDLIFE SERVICE. 2010. Prairie Pothole Region, Smart Science. Mountain Prairie Region. http://www.fws.gov/mountain-prairie/factsheets/Smart%20Science %20Tip%20Sheet.pdf Accessed Feb. 2, 2012. VERMEIRE, L. T. AND M. J. RINELLA. 2009. Fire alters emergence of invasive plant species from soil surface deposited seeds. Weed Sci., 57:304–310. ——— AND A. D. ROTH. 2011. Plains Prickly Pear response to fire: effects of fuelload, heat, fireweather, and donor site soil. Rangeland Ecol. Manage., 64:404–413. WHELAN, R. J. 1995. The ecology of fire. Cambridge University Press, Cambridge, U.K. 346 p. WHISENANT, S. G. 1990. Postfire population dynamics of Bromus japonicas. Am. Midl. Nat., 123:301–308. WILLSON, G. D. AND J. STUBBENDIECK. 1997. Fire effects on four growth stages of smooth brome (Bromus inermis Leyss.). Nat. Area J., 17:306–312. WOTTON, M. 2009. Grass fire behaviour and flame. http://www.firelab.utoronto.ca/behaviour/grass_ fire.html. Accessed December 2013. SUBMITTED 7 SEPTEMBER 2013 ACCEPTED 27 SEPTEMBER 2014