This article was downloaded by: [Matt Madsen] #742 On: 17 September 2012, At: 14:25 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Arid Land Research and Management Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/uasr20 Influence of Soil Water Repellency on Seedling Emergence and Plant Survival in a Burned Semi-Arid Woodland a b b Matthew D. Madsen , Steven L. Petersen , Kaitlynn J. Fernelius , b c Bruce A. Roundy , Alan G. Taylor & Bryan G. Hopkins b a USDA–Agricultural Research Service, Eastern Oregon Agricultural Research Center, Burns, Oregon, USA b Department of Plant and Wildlife Sciences, Brigham Young University, Provo, Utah, USA c Department of Horticulture, Cornell University, Geneva, New York, USA To cite this article: Matthew D. Madsen, Steven L. Petersen, Kaitlynn J. Fernelius, Bruce A. Roundy, Alan G. Taylor & Bryan G. Hopkins (2012): Influence of Soil Water Repellency on Seedling Emergence and Plant Survival in a Burned Semi-Arid Woodland, Arid Land Research and Management, 26:3, 236-249 To link to this article: http://dx.doi.org/10.1080/15324982.2012.680655 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. #742 Arid Land Research and Management, 26:236–249, 2012 Copyright # Taylor & Francis Group, LLC ISSN: 1532-4982 print=1532-4990 online DOI: 10.1080/15324982.2012.680655 Influence of Soil Water Repellency on Seedling Emergence and Plant Survival in a Burned Semi-Arid Woodland Matthew D. Madsen1, Steven L. Petersen2, Kaitlynn J. Fernelius2, Bruce A. Roundy2, Alan G. Taylor3, and Bryan G. Hopkins2 Downloaded by [Matt Madsen] at 14:25 17 September 2012 1 USDA–Agricultural Research Service, Eastern Oregon Agricultural Research Center, Burns, Oregon, USA 2 Department of Plant and Wildlife Sciences, Brigham Young University, Provo, Utah, USA 3 Department of Horticulture, Cornell University, Geneva, New York, USA High intensity wildfires in semiarid shrub and woodland plant communities can leave ecosystems incapable of self-repair and susceptible to weed invasion. Subsequently, land managers need effective restoration tools to reseed native vegetation back into these degraded systems. In order to develop successful post-fire restoration approaches in these communities, it is critical that we understand the mechanisms that impair reseeding success. Our objective was to quantify the influence of soil water repellency on seedling emergence and plant growth in a greenhouse study using soil cores obtained from beneath burned Juniperus osteosperma trees. Soil cores were seeded with either Elymus wawawaiensis or Agropyron cristatum, and watered with either a high (watered daily) or a low water regime (watered every 5 days). During the first watering event, water repellency was ameliorated in half the cores by adding a wetting-agent comprised of alkylpolyglycoside-ethylene oxide=propylene oxide block copolymers. Results showed that water repellency reduced seedling emergence and seedling survival by decreasing soil moisture availability. Wetting-agents improved ecohydrologic properties required for plant growth by decreasing runoff and increasing the amount and duration of available water for seedling emergence, survival, and plant growth. These results indicate that soil water repellency can act as an ecological threshold by impairing establishment of reseeded species after a fire. Where restoration efforts are limited by soil water repellency, Received 18 April 2011; accepted 30 November 2011. We are grateful to Daniel Zvirzdin and Alexander Zvirzdin for their aid in conducting this research. Wetting agent and technical support was provided by Stan Kostka (Aquatrols, Paulsboro, NJ, USA). Funding was provided by the USDA-Natural Resource Conservation Service, Conservation Initiative Grant, USDA-National Institute of Food and Agriculture’s Rangeland Research Program, and the USDA-Agricultural Research Service. Mention of a proprietary product does not constitute a guarantee or warranty of the product by USDA or the authors and does not imply its approval to the exclusion of the other products that also may be suitable. USDA is an equal opportunity provider and employer. Address correspondence to Matthew D. Madsen, USDA–Agricultural Research Service 67826-A Hwy 205, Eastern Oregon Agricultural Research Center, Burns, OR 97720, USA. E-mail: matthew.madsen@oregonstate.edu 236 Wildfire Restoration in Water Repellent Soil 237 wetting agents have the potential to improve the success of post-fire reseeding efforts. Future work is needed to validate these findings in the field. Downloaded by [Matt Madsen] at 14:25 17 September 2012 Keywords erosion, hydrophobicity, pinyon-juniper, reseeding, restoration, revegetation, surfactants, weeds, wetting-agents, wildfire High intensity or catastrophic wildfires in semiarid shrub and woodland plant communities can leave these ecosystems incapable of self-repair and susceptible to weed invasion (Young & Evans, 1978; Brooks et al., 2004). Invasive species such as Bromus tectorum L. (cheatgrass) have a negative impact on rangeland ecological services, including decreased forage production, watershed stability, and water quality (Young & Evans, 1978; Miller & Tausch, 2001). These flammable weeds also alter the frequency, intensity, extent, and seasonality of natural fires, thus further promoting weed dominance and loss of native species (D’Antonio & Vitousek, 1992). One approach to preclude weed invasion is to reseed the site with desirable species (Epanchin-Niell, Englin, & Nalle, 2009). Through this approach, recovery is aided by replenishing depleted seed pools, which helps restore ecosystem processes while reducing resource availability for invasive weeds (Cox & Anderson, 2004). In the past, land managers typically selected introduced species for post-fire restoration, such as Agropyron cristatum (L.) Gaertn. (crested wheatgrass), and Thinopyrum intermedium (Host) Barkworth & D.R (intermediate wheatgrass). These species can increase establishment success, lower seeding costs, provide better weed competition, and improve livestock forage quality (Steward, 1950). Currently, fire rehabilitation programs are increasing the use of native plant materials in place of introduced species in an effort to reinstate ecosystem processes and improve species diversity after a fire (Thompson et al., 2006). However, these native species are costly and establishment success is typically less than desirable (Roundy et al., 1997). Consequently, land managers throughout the Intermountain West are requesting new techniques that will improve establishment success of native plant materials in order to restore habitats lost to wildfire, prevent subsequent weed dominance, and restore natural fire cycles. In order to develop successful post-fire restoration approaches, it is critical to understand the factors that impair native vegetation recovery. Soil water repellency (or hydrophobicity), is one factor that may significantly limit post-fire recovery in Pinus-Juniperus systems and promote subsequent weed domination. Soil water repellency is commonly found in arid and semi-arid ecosystems in unburned soils (Krammes & DeBano, 1965; Doerr et al., 2000; Jaramillo et al., 2000). Sites with coarse-textured soils that contain woody evergreen plants, which have high amounts of resins, waxes, or aromatic oils, and associated thick litter layers, are most susceptible to soil water repellency development (Doerr et al., 2000; Jaramillo et al., 2000). However, it cannot be assumed that all sites that have these soil and plant characteristics will be associated with soil water repellency, as there are multiple complex processes associated with water repellency formation (Doerr et al., 2009). In unburned conditions soil water repellency can have a strong control on precipitation inputs. During a precipitation event, thick litter layers can help retain precipitation from running off site, while underlying water repellent soil concentrates and directs this moisture through preferential flow channels to greater depths than could be achieved on wettable soil (Madsen et al., 2008; Robinson et al., 2010). In Downloaded by [Matt Madsen] at 14:25 17 September 2012 238 M. D. Madsen et al. water limiting environments, soil water repellency may benefit woody plants by minimizing near surface evaporation and increasing soil moisture within the root zone (Madsen et al., 2008; Robinson et al., 2010). Burning can alter the severity and depth of the water repellent layer as heat volatilizes organic substances in the litter and upper water repellent soil layers (DeBano and Krammes, 1966). These molecules penetrate into the cool underlying soil layers where they condense around soil particles and either render them hydrophobic or further decrease their affinity for water (Savage, 1974; Doerr et al., 2000). Pinus spp. (piñon) and Juniperus spp. (juniper) are examples of woody vegetation types which under both unburned and burned conditions can be strongly correlated with the presence of soil water repellency (Jaramillo et al., 2000; Madsen et al., 2008; Pierson et al., 2009). In the western United States, Pinus-Juniperus woodlands have expanded since Eurasian settlement in the late nineteenth century, into primarily sagebrush steppe and grassland communities (Burkhardt & Tisdale, 1976). These species out-compete understory species, which decreases the potential for frequent, moderate-severity fires carried through the understory vegetation, and increases the potential for less-frequent large-scale catastrophic crown fires (Miller & Tausch, 2001). Madsen et al. (2011) demonstrated that within a Pinus-Juniperus woodland, the spatial distribution of water repellency was strongly correlated with pre-fire tree canopy cover. In this study critical water repellency (i.e., water repellency that can be measured through the water drop penetration test (WDPT) (Krammes and DeBano, 1965) extended from the base of the tree to just beyond the canopy edge. Subcritical water repellency (i.e., water repellency levels are insufficient to be detected through the WDPT test but are at sufficient levels to influence soil infiltration; Tillman et al., 1989) extended half a canopy radius past the edge of the critical water repellency zone. At sites where critical water repellency was present, vegetation cover and density were dramatically reduced relative to non-water repellent sites; these variables were also reduced in soils with subcritical water repellency, albeit to a lesser extent. These results by Madsen et al. (2011) and other authors, such as the research of Adams et al. (1970) on Larrea tridentata (creosote bush) and the studies of Salih et al. (1971) with Artemisia tridentata (sagebrush), may indicate that soil water repellency limits revegetation success immediately following fire. Consequently, restoration approaches that focus on treating soil water repellency issues could potentially improve the success of seeding projects while simultaneously decreasing runoff and soil erosion, and preventing weed invasion. Use of commercially available surface active agents (wetting-agents or surfactants) may provide an alternative post-fire restoration approach where water repellency inhibits site recovery. A wide variety of ionic and nonionic wetting-agents are produced commercially, ranging from simple dish soaps to sophisticated polymers chemically engineered to overcome soil water repellency. Wetting-agents are generally organic molecules that are amphiphilic (hydrophobic tails and hydrophilic heads). While wetting-agents have different modes of action, water repellent soils are rendered wettable by the hydrophobic tail of the wetting-agent chemically bonding to the non-polar hydrophobic coating on the soil particle, while the hydrophilic head of the molecule attracts water molecules into the soil (Cisar et al. 2000; Kostka 2000). Various small plot post-fire research projects located in the mountains of southern California revealed that the application of wetting-agents after a fire can reduce soil erosion and improve vegetation establishment (Osborn et al., 1967; Krammes & Osborn, 1969; Debano & Conrad, 1974). These studies Downloaded by [Matt Madsen] at 14:25 17 September 2012 Wildfire Restoration in Water Repellent Soil 239 suggest that wetting-agent applications can be a successful post-fire treatment. While wetting-agents have not been used in wildland systems since the 1970’s, they were extensively used and further developed within various aspects of the agricultural industry, with most applications in turfgrass systems (Cisar et al., 2000; Kostka & Bially, 2005). These newly developed wetting-agents may provide an innovative approach for alleviating the effects of water repellency on the establishment of seeded species. The primary objective of this research was to quantify the extent that soil water repellency within burned Pinus-Juniperus woodland soils influences emergence and growth of the non-native bunchgrass A.cristatum, and native bunchgrass, Elymus wawawaiensis J. Carlson and Barkworth (Snake River wheatgrass), and in conjunction with this objective, determine if wetting-agent can be used to reduce soil water repellency and subsequently improve seedling emergence and survival. We hypothesized that soil water repellency would decrease seedling survival by decreasing soil moisture availability, and that the reduction of soil water repellency through wetting-agent application would overcome these limitations. Methods Soil Collection Research was conducted on soil obtained from the boundaries of Utah’s largest wildfire on record (145,000 ha), the 2007 Milford Flat wildfire. Soils were collected approximately 13.7 km NW of Milford, UT, USA (Lat: 38 26’ 12’’ N, Long: 112 51’ 46’’ W, elevation 1847 m). A detailed description of the site is given by Madsen et al. (2011). The site is predominantly on west-facing slopes, positioned at the base of the Mineral Mountain Range. Parent material is characterized as alluvium derived from acid and intermediate igneous rock, with soil classified as coarse sandy loam, mixed, mesic Aridic Haploxerolls (3–10% slope). Volumetric soil water content at 1.5 MPa (permanent wilting point) and 0.33 MPa (field capacity) was equal to 12.8%, and 24.8%, respectively (Soil Survey Staff, 2011). Mean annual precipitation is approximately 370 mm (PRISM Climate Group, 2009). Prior to the fire, the plant community was a Phase III, Pinus-Juniperus woodland, with Juniperus osteosperma (Torr.) Little (Utah juniper), and Pinus monophylla Torr. & Frém. (singleleaf piñon) trees acting as the primary plant layer driving ecological processes (Miller et al., 2008). Soil was collected one year after the fire, from the subcanopy of burned J. osteosperma trees. At the time of collection, the soil remained almost completely bare of live vegetation, with trace amounts of annual forbs and grasses (3.1 1.1 plants m2) (average and standard error of the mean). The upper layer of the soil was wettable and appeared to be primarily ash originating from burned litter and debris; this layer extended from the soil surface down to 1.7 0.1 cm. Below this depth (1.7–4.5 cm), water repellent mineral soil predominated. Average WDPT; within the water repellent layer was 5,256 258 sec. Soil cores were obtained by pressing polyvinyl chloride (PVC) tubes (20.3 cm diameter by 36-cm deep) into the soil with the bucket of a front-end loader. To minimize soil compaction, the end of each tube was sharpened before being pressed into the soil. Cores were modified for use as growing pots by fastening a woven porous landscape fabric to the bottom of the PVC pipe. To allow for drainage at the soil surface, pots were fitted with a runoff spout 19.1 mm in diameter (Figure 1). Pebbles were placed in front of the spout to minimize seed and soil loss. Downloaded by [Matt Madsen] at 14:25 17 September 2012 240 M. D. Madsen et al. Figure 1. A. Diagram of rainfall simulator; B. soil core modified for use as a growing pot, attached with a runoff catchment tube. Study Design This experiment was conducted in a greenhouse located on the campus of Brigham Young University (BYU), Provo UT, USA. Treatments included soil cores treated with and without wetting-agent, two watering regimes, and two seeded species, with all combinations arranged in a randomized block design, with three soil core replicates per block, and five blocks in the study (2 soil treatments 2 watering regimes 2 species 3 core-replicates 5 blocks ¼ 120 soil cores in the study). Each pot was seeded with 35 seeds of either A. cristatum or E. wawawaiensis. Total germination of A. cristatum and E. wawawaiensis, was 90% and 85%, respectively. During the first watering event, we applied the non-ionic wetting-agent IrrigAid Gold (Aquatrols, Paulsboro, NJ, USA), which is composed of a blend of alkylpolyglycoside and ethylene oxide=propylene oxide block copolymers, at 0.012 ml cm2. In preliminary tests, we found that this wetting-agent concentration was required to allow complete wetting of the soil column. After the pots were treated with wetting-agent, we verified that the soil had been made wettable using the WDPT test. As Pinus-Juniperus woodlands typically occur on sloping terrain and the influence of soil water repellency can be more pronounced on a slope, pots were placed on a 15 angled bench, similar to methods described by Osborn et al. (1967). We watered the pots by running a rainfall simulator with the bottom of the simulator 40 cm above the soil surface (Figure 1). Individual rainfall simulators were designed to water one pot at a time, which were constructed from seven liter buckets that had a 25.4 cm radius bottom, and 50 drip tubes (10 mm long, with an inside diameter of 0.864 mm and wall thickness of 0.305 mm), to distribute the water. The rate of flow Wildfire Restoration in Water Repellent Soil 241 Downloaded by [Matt Madsen] at 14:25 17 September 2012 from the bucket was controlled by adjusting the head pressure using a Mariotte tube (4.8 mm diameter) that slid within a bored rubber stopper attached to the top of the bucket. To start and stop the flow of water from the rainfall simulator, surgical tubing and a clamp were placed at the top of the Mariotte tube. Pots were watered under either a low or a high watering regime. The low watering regime was designed to replicate spring-like conditions, with periods of adequate soil moisture, followed by intermittent periods of no precipitation. This watering treatment consisted of watering daily for the first 4 days, to insure seed germination, then once every 5 days. We also watered pots daily on days 20–24 to see if seeds not yet germinated would emerge. The high water regime was designed so the seedlings would not be water stressed. Under this watering regime, we watered the pots daily throughout the project. At each watering (low and high) we delivered a total of 0.62 cm (400 mL) of water at a rate of 3.0 cm hr1. Measurements The experiment was conducted for 48 days from the time of seeding (14 September 2008–2 November 2008). During this period, runoff was collected from three randomly-chosen blocks. Runoff was measured from each pot within a catchment tube made of 4.5-cm-diameter thin-wall PVC cylinders, with the cap hung just below the runoff spout of each pot (Figure 1). For each different watering regime and soil treatment, soil water content was measured on four randomly-selected pots. Soil water content was continuously recorded, at the 1–3 cm soil depth using EC-5 Soil Moisture Sensors in conjunction with Em5b data loggers (Decagon Devices, Pullman, WA, USA). For all blocks, the number of live seedlings was recorded every 2 days throughout the experiment. Upon harvest, below-ground and above-ground biomass was measured separately after drying at 65 C for 72 h. Data Analysis Mixed model analysis was used to analyze runoff, soil water content, final seedling density, above-ground biomass, and below-ground biomass (SYSTAT 12, Systat Software Inc. Chicago, IL, USA; P < 0.05). Blocks were considered random, while species, watering regimes, and soil treatments were considered fixed factors. Runoff and soil water content were analyzed as repeated measures. For treatments found to affect the response variables, mean values were separated using Tukey’s Honestly-SignificantDifference test. Pairwise comparison among the treatments for runoff and soil water content response variables were averaged by treatment over the period of the study. Analysis of soil water content was performed on values at the time of watering (i.e., peak SWC), and 4 days after watering (i.e., trough SWC). Final seed density was analyzed by the number of live plants at the end of the study and after normalizing the density data, by species, with the percent of germinable seeds planted. Results Runoff Runoff was not influenced by species, or watering regime, but was influenced by wetting-agent treatment (Table 1A). The control had, on average, 24%, and 19% of the added water runoff, while the wetting-agent treatment decreased runoff with 242 M. D. Madsen et al. Table 1. A. Mixed-model analysis results indicating significance of water regime and wetting-agent treatment on runoff and soil water content just after watering (SWC: peak) and four days after watering (SWC: trough). B. Pair-wise comparisons showing mean and associated standard error for the response variables that were found to be significant Hydrologic Response A. Response variables Runoff Downloaded by [Matt Madsen] at 14:25 17 September 2012 Effect variables Water regime Wetting-agent Species SWC: trough F P F P F P 2.3 178.3 0.002 0.144 <0.001 0.966 54.5 8.9 – <0.001 0.004 – 139.1 17.1 – <0.001 0.001 – Pair-wise comparisons B. Runoff (mL) low water þ control low water þ WA high water þ control high water þ WA SWCa: peak 94.7 5.7 18.0 3.4 76.1 5.8 22.1 3.6 a b a b SWC: peak (%) 17 2 25 2 31 1 33 1 a b c c SWC: trough (%) 62 15 2 27 1 30 1 a b c c Means without a common letter differ at P < 0.05, Tukey HSD test. a SWC ¼ soil water content. only 4%, and 5% of the water runoff, for the low and high watering regimes, respectively (Table 1). At harvest, it was observed that the majority of the control pots still had a band of air-dry water-repellent soil just below the soil surface (Figure 2). It was speculated that for the control pots, water that did not run off from the soil cores infiltrated around the water repellent layer along the sides of the pot, into the wettable soil below. In contrast, wetting-agent treated soils appeared to allow for an even wetting of the entire soil core (Figure 2). Soil Water Content Soil water content was affected by watering regime and wetting-agent treatment (Table 1A). Under the low watering regime, soil water content was higher in the wetting-agent treated soil for both the peak and trough periods in relationship to the control (Table 1B). Under the high watering regime, soil water content was statistically similar, although the peaks and troughs on average were higher for wetting-agent treated soil (Table 1B). Density Seedling density was influenced by watering regime, wetting-agent application, and species (Table 2). However, relative seedling density, normalized by percent of Downloaded by [Matt Madsen] at 14:25 17 September 2012 Wildfire Restoration in Water Repellent Soil 243 Figure 2. Aerial view and cross section photo of E. wawawaiensis seedlings grown under a low water regime on: A. water repellent soil; B. water repellent soil that was treated with wetting-agent (WA). germinable seeds, did not differ by species (Table 2). Under the low water regime, within the control pots, seedling density first peaked at about 10 days after seeding (Figure 3). Under the same watering regime, wetting-agent treated pots peaked at day 14, with seedling densities 268% higher than the control peak densities. Between days 10–20, 72% of the seedlings growing in the control desiccated (Figure 3). Unlike the control pots, wetting-agent treated pots under the low water regime had relatively few plants desiccate before the next daily watering period, with a drop in seedling density of only 9%. Daily watering on days 20–24 produced a minor increase in seedling densities for control and wetting-agent treatments, as a result of seeds emerging that had not Table 2. Mixed-model analysis of Agropyron cristatum and Elymus wawawaiensis seedling responses to water regime, and wetting-agent treatment Vegetation response Effect variables Water regime Response variables Seedling survival Survival of germinable seeds Above-ground biomass Below-ground biomass Wetting agent Species F P F P F P 129.1 130.2 100.1 82.6 <0.001 <0.001 <0.001 <0.001 102.1 102.0 50.1 59.7 <0.001 <0.001 <0.001 <0.001 4.9 1.8 11.3 2.4 0.034 0.191 0.002 0.128 Downloaded by [Matt Madsen] at 14:25 17 September 2012 244 M. D. Madsen et al. Figure 3. Combined results for A. cristatum and E. wawawaiensis seedling densities over the course of the experiment, with results normalized by total germination for pots watered under a low or high watering regime, treated with or without wetting-agent (WA). Significant differences (P < 0.05) at the end of the experiment are shown by unique letters. emerged previously under the low water regime (Figure 3). At the end of the experiment under the low watering regime, wetting-agent treated pots had significantly higher final seedling densities, with 292% more seedlings in the wetting agent treatment in comparison to the control (i.e., 12.2 and 46.3% of the germinable seeds planted produced a live plant at the end of the study for control and wetting-agent treatment, respectively). Under the high water regime, control pot seedling densities steadily increased over the course of the experiment, with seedling densities similar to wetting-agent treated soils under the low watering regime (Figure 3). Wetting-agent treated pots under the high regime also had higher seedling densities than the control pots, with 49.6% higher seedling survival (i.e., 50.2 and 74.9% of the germinable seeds planted produced a live plant at the end of the study for control and wettingagent treated pots, respectively). Biomass Total above-ground biomass was influenced by watering regime, wetting-agent application, and species (Table 2). Pots treated with wetting-agent under the low water regime had above-ground biomass 690% and 722% greater than the control, for A. cristatum and E. wawawaiensis respectively (Figure 4). Above-ground biomass of wetting-agent treated pots for both species, under the low watering regime was statistically similar to the control under the high watering regime. Under the high watering regime, biomass of seedlings in the wetting-agent treated pots was statistically higher than seedlings in the control, with above-ground biomass 50% and 66% greater than the control for A. cristatum and E. wawawaiensis, respectively (Figure 4). However, under the high watering regime biomass of A. cristatum was similar to that of E. wawawaiensis grown in wetting-agent treated soil. Below-ground biomass was influenced by watering regime and wetting-agent application, but unlike above-ground biomass it did not differ by species (Table 2). Of all the response variables measured in this study, below-ground biomass showed Downloaded by [Matt Madsen] at 14:25 17 September 2012 Wildfire Restoration in Water Repellent Soil 245 Figure 4. Above and below ground biomass of A. cristatum and E. wawawaiensis grown on water-repellent soil treated with or without wetting-agent (WA), under high and low watering regimes. Significant differences (P < 0.05) at the end of the experiment are shown by unique letters. the greatest increase with wetting-agent treatment. Pots treated with wetting-agent under the low water regime had below-ground biomass 1056% and 1196%, greater than the control, for A. cristatum and E. wawawaiensis, respectively. Similar to above-ground biomass, wetting-agent treated pots under the low watering regime had similar above-ground biomass as control treatments under the high watering regime, but less biomass than wetting-agent treated pots under the higher watering regime. Under the high water regime, above-ground biomass in the wettingagent pots was greater than the control by 119%, and 127%, for A. cristatum and E. wawawaiensis, respectively. While harvesting the control pots, we observed that seedling roots generally did not grow through the water repellent layer, but grew along the sides of the soil pot to avoid the water repellent layer. In contrast, roots were evenly distributed throughout the wetting-agent treated soil (Figure 2). Discussion These results are consistent with previous wildland fire studies that have documented decreased germination and seedling survival as a result of soil water repellency Downloaded by [Matt Madsen] at 14:25 17 September 2012 246 M. D. Madsen et al. (DeBano, 1969; Krammes & Osborn, 1969; DeBano & Conrad, 1974), and this is the first study to demonstrate that water repellency limits reseeding success within Pinus-Juniperus woodland soils. Madsen et al. (2011) examined the spatial distribution and severity of soil water repellency within the area the soil was collected for this study, and determined that revegetation success was correlated to these parameters. However, the methods implied by Madsen et al. (2011) do not imply causation. In this study causation was shown by using wetting-agent to cancel out (or ameliorate) the effect of soil water repellency and allow us to compare revegetation on water repellent and non-water repellent soils. Specifically the results of this greenhouse experiment suggest that within burned Pinus-Juniperus woodlands, soil water repellency can promote runoff, decrease soil water content, and impair seedling emergence, growth, and plant survival. In this study decreased soil water content near the soil surface is most likely associated with soil water repellency promoting runoff (Table 1) and directing infiltrating water through preferential flow channels below the water repellent soil (Dekker & Ritsema, 2000). Consequently, water repellency may impair seeding success by decreasing moisture availability to levels that are not adequate for seed germination and seedling survival. Potentially the cycle of seedling emergence and desiccation that was observed in this study may also occur in the field, which would result in a loss of residual or aerially-seeded species from the seed bank. Once soil water repellency has dissipated after a fire, and seed propagules of desired species are no longer present, the site may be primed for weed invasion. In general, the susceptibility of a site to weed invasion can be explained in part, by the fluctuating resource hypothesis which states that ‘‘a plant community becomes more susceptible to invasion whenever there is an increase in the amount of unused resources’’ (Davis et al., 2000, p. 529). Therefore, we hypothesize that soil water repellency negatively affects germination of seeded species and increases invasibility by annual weeds, such as B. tectorum, that can establish on the resource rich subcanopy areas after soil water repellency dissipates. Results of this study are limited to the specific site in which soil was obtained from the study (i.e., subcanopy region within a phase III Pinus-Juniperus woodland). The variables that determine the occurrence, severity, persistence, and spatial distribution of water repellency across the landscape are likely associated with various ecological attributes such as soil properties, topography, and woodland stand characteristics (Lewis et al., 2006). For areas where soil water repellency is limiting vegetation recovery, these results indicate that wetting-agents can ameliorate post-fire water repellent soil and subsequently help restore ecohydrologic function in conjunction with reseeding efforts. Application of wetting-agents in this study significantly decreased water runoff and increased the amount of available soil moisture, which increased plant density, and above and below ground biomass. These results are also consistent with previous studies within the California chaparral which showed that wetting-agents improve seedling densities on post-fire water repellent soil (e.g., Osborn et al., 1967; Krammes & Osborn, 1969; Debano & Conrad, 1974). This research, along with previous studies in other ecosystems, provides evidence that this technology should be further field-tested in wildland systems impaired by post-fire water repellent soil. Conclusions and Management Implications The effects of large, high-severity wildfires are a significant management concern for rangeland ecosystems. After a fire, the ability of an ecosystem to recover is Downloaded by [Matt Madsen] at 14:25 17 September 2012 Wildfire Restoration in Water Repellent Soil 247 dependent on the extent to which ecological processes have been altered, both prior to and as a result of the fire. This study indicates that the modification of the soil through the development of a post-fire water repellent layer is one alteration that may significantly limit site recovery. Restoration approaches that focus on ameliorating water repellency could potentially improve the success of native plant materials in post-fire reseeding efforts. This study provides evidence that the use of wetting-agents can significantly improve ecohydrologic properties required for plant growth within a post-fire Pinus-Juniperus woodland. While these results are promising, future work is needed to verify these results in the field, and to determine if the application of this soil amendment would be practical for post-fire wildland revegetation. The application of soil amendments is typically not practical for the reclamation of rangeland systems due to large areas and difficult topography that require treatment, as well as the perceived low economic value of the land. 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