Lake Tahoe Upland Fuels Research Project: Investigating the effects of fuels reduction treatments on forest structure, fire risk, and wildlife Final Report to U.S. Department of the Interior Bureau of Land Management November 2, 2012 Principal Investigators: Patricia N. Manley, Ph.D. a b Dennis D. Murphy, Ph.D. c Bruce M. Pavlik, Ph.D. Research Team: Alison E. Stanton c Traynor G. Biasiolli a Angela M. White, Ph.D. a Matthew D. Timmer a Brandon M. Collins, Ph.D. a Gary B. Roller a a U.S. Forest Service, Pacific Southwest Research Station, Davis, CA b Department of Biology, University of Nevada, Reno, NV c BMP Ecosciences, 156 South Park, San Francisco, CA 1 TABLE OF CONTENTS MANAGEMENT SUMMARY .................................................................................................................. 3 1.0 INTRODUCTION AND BACKGROUND ......................................................................................... 4 2.0 GOALS AND OBJECTIVES............................................................................................................... 5 2.1 CURRENT STATE OF KNOWLEDGE ................................................................................................... 5 2.2 PROJECT OBJECTIVES ....................................................................................................................... 6 3.0 METHODS ............................................................................................................................................ 6 3.1 STUDY SITES ....................................................................................................................................... 6 3.2 FUELS REDUCTION TREATMENTS .................................................................................................. 11 3.3 SAMPLING DESIGN AND TIMELINE ................................................................................................. 12 3.4 VEGETATION AND FUELS ................................................................................................................ 14 3.5 WILDLIFE ......................................................................................................................................... 14 3.6 DATA ANALYSIS ............................................................................................................................... 15 3.6.1 Vegetation and Fuels ............................................................................................................... 15 3.6.2 Wildlife...................................................................................................................................... 16 4.0 RESULTS ............................................................................................................................................ 18 4.1 VEGETATION .................................................................................................................................... 18 4.2 FUELS ................................................................................................................................................ 24 4.3 SMALL MAMMALS ........................................................................................................................... 29 4.4 BIRDS ................................................................................................................................................ 31 4.5 INVERTEBRATES............................................................................................................................... 34 5.0 DISCUSSION ...................................................................................................................................... 36 5.1 VEGETATION AND FUELS ................................................................................................................ 36 5.2 WILDLIFE ......................................................................................................................................... 37 6.0 CONCLUSION ................................................................................................................................... 39 LITERATURE CITED ............................................................................................................................ 41 APPENDICES ........................................................................................................................................... 44 2 MANAGEMENT SUMMARY BACKGROUND • • • • The National Fire Plan and the Healthy Forest Restoration Act mandate federal land managers to restore forest habitats and reduce the risk of wildfire, particularly in the wildland-urban interface. The Healthy Forest Initiative implementation plan includes monitoring and tracking performance to confirm that these objectives are being met. This report summarizes the results of a comprehensive six year experiment investigating the effects of fuels reduction treatments on fuel conditions and fire risk, forest structure, and wildlife in the Lake Tahoe Basin. Data were collected for eight paired treatment and control units (n=16) on the west and east shores of the Basin. Three of the west shore units were in protected activity centers (PACs) for California Spotted Owl (Strix occidentalis occidentalis) and Northern Goshawk (Accipiter gentilis). We collected detailed data on vegetation structure and composition, fuel loading, birds, small mammals, and ants before and after treatments were implemented. KEY RESULTS • • • • • • Mechanical fuels reduction treatments resulted in more open forest with fewer smaller diameter trees. Post-treatment tree species composition was less fir-dominated and included a greater proportion of pines. In the understory, mechanical treatments reduced shrub cover and coarse woody debris but did not affect herbaceous cover. Hand thinning reduced coarse woody debris, but not vegetative cover. Treatments successfully reduced both proportion of area killed and large tree mortality under modeled fire conditions. There is little evidence to support the removal of large proportions of basal area for fuel reduction. There are other valid justifications for large basal area reductions, e.g., restoration, economics, but fuel reduction is not one. Small mammal community richness and evenness shifted negligibly following treatment. Overall abundance of mammals increased non-significantly following treatment. Measures of bird community richness and evenness were largely unchanged following treatment. Pooled abundance of all bird species declined slightly after treatment. Ant community richness did not differ between pre- and post-treatment forests. We found no differences in site occupancy for any species following treatment implementation. IMPLICATIONS FOR MANAGEMENT Our results indicate that implementation of conventional fuels reduction treatments is consistent with maintenance of bird, small mammal, and ant community diversity over short time frames. Fuels treatments met fire risk objectives while resulting in markedly similar post-treatment wildlife communities. Importantly, reduction of surface fuels resulted in greater tree resiliency to future wildfires over the lifespan of these treatments. While wildlife communities were largely unchanged, we observed substantial changes in abundance of several species following treatments, as well as a slight increase in overall mammal abundance and a slight decline in bird abundance. Further research is needed to investigate longer-term impacts of fuels treatments on wildlife communities. 3 1.0 INTRODUCTION AND BACKGROUND In response to the elevated threat of high intensity wildfire throughout much of the western United States, the National Fire Plan and the Healthy Forest Restoration Act (HFRA) mandated federal land managers to restore forest habitats and reduce the risk of wildfire, particularly in the wildland-urban interface (WUI). In general, these policies set a goal of establishing pre-European conditions or a historic range of variability that predated modern fire suppression (Parsons et al. 1999, Stephenson 1999). Restoration may employ an integrated strategy that includes tree thinning and brush removal, collectively called “fuels treatments.” The principal goals of fuels reduction treatments are (1) to reduce the amount of burnable materials, thereby reducing fire intensities and crown fire potential, and improving suppression capability (Agee 2002), and (2) to improve forest health and ecological integrity. The restoration of the natural fire regime, if possible, is likely to be the most effective approach to achieving desired conditions for forests in the Basin. However, many obstacles limit managers’ ability to implement a natural fire regime. A significant portion of the Lake Tahoe Basin is considered a high-risk environment for severe wildfires (Murphy et al. 2006). This elevated wildfire threat is a result of land use practices over the last 150 years. In the 1860’s extensive logging resulted in the development of single-aged stands. By the turn of the 20th century, nearly two-thirds of the entire Basin had been clear-cut. The clearing was concentrated in lower elevation pine forests, with more than 60% of this forest type removed (Murphy and Knopp 2000). Recovery of the forest over the last century was altered by management focused on fire suppression. Recently published research suggests that current fire return intervals in the Lake Tahoe Basin are at their lowest levels in at least the last 14,000 years (Beaty and Taylor 2009). Modern forests that developed under fire suppression after extensive logging are overly dense and crowded with small trees and contain an unnaturally high level of fuels (Barbour et al. 2002, Taylor 2004). Addressing the wildfire threat and implementing restoration measures in the Lake Tahoe Basin presents unique challenges. First, the topography and climate in the Basin, in combination with clear-cutting and fire exclusion, have produced fir- (Abies sp.) dominated forests in the lower montane zone, thus increasing ladder fuels and probability of crown fires. Second, much of the fuel load in the Basin is in the form of small trees with low commercial value, thus reducing the cost effectiveness of mechanical biomass removal. Third, the proximity of these forests to populated areas and the importance of tourism have limited the use of prescribed burning as a management option. Smoke and liability issues, along with a small number of allotted burn days in many years has severely limited the number of acres that have been treated with fire. Despite these challenges, fire risk reduction has become a top priority in the Basin and funding to implement fuels reduction treatments has increased in recent years. The National Fire Plan, the HFRA, and the Sierra Nevada Public Lands Management Act (SNPLMA) have made more funds available. In addition, the Lake Tahoe Basin Management Unit (hereafter LTBMU) has continued treatments on more than 12,000 acres (in defense and threat zones) as part of the South Shore Fuels Reduction and Healthy Forest Project. Project Coordination The Lake Tahoe Upland Fuels Research Project science team represents collaboration between the US Forest Service Pacific Southwest Research Station (PSW), University of Nevada, Reno (UNR), and BMP 4 Ecosciences (a research consultant). The principle investigators in this project are Patricia Manley (PSW), Dennis Murphy (UNR), and Bruce Pavlik (BMP). This team collaborated closely with the LTBMU to identify available treatment units. The Upland Fuels Research Project (UPFU) received SNPLMA funds under the Environmental Improvement Program (EIP#10123) to evaluate vegetation composition, structure, fuel loading, and animal communities (vertebrate and invertebrate) on six sites on the west shore slated for fuels reduction and in six paired, untreated controls on adjacent federal, private, and state lands. Documents summarizing results for 2006 were prepared (Stanton and Dailey 2007)Stanton and Dailey 2007). A grant from Nevada State License Plate Funds (LTLP 07-01) funded a second year of sampling in 2007 on the west-shore sites and established two new pairs of sites on the east shore on Nevada State Lands. Two additional grants from Nevada State License Plate Funds (LTLP 08-10 and 0902) completed sampling and analysis for the two pairs of east-side sites, respectively. This report summarizes the results of a comprehensive six year experiment that evaluated the effects of fuels reduction treatments on fuel conditions and fire risk, forest structure, and wildlife in the LTBMU. 2.0 GOALS AND OBJECTIVES 2.1 CURRENT STATE OF KNOWLEDGE A large body of scientific literature addresses many issues associated with wildland fire and different fuel treatments applied in various vegetation types. Substantial portions of that work primarily address the use of prescribed fire. Evidence supports the utility of prescribed fire in reducing crown-fire potential or improving the resilience of forest stands to wildfire, but these studies are largely based on informal observations (Brown 2002, Carey and Schumann 2003), post-fire inference (Omi and Kalabokidis 1991, Pollet and Omi 2002) and modeling (Stephens 1998, Finney 2001, Agee and Skinner 2005, Peterson et al. 2005). Limited scientific consensus exists regarding the specifics of how treatments are implemented and the relative effectiveness of different prescriptions across vegetation types (Carey and Schumann 2003, Peterson et al. 2005, Stephens and Ruth 2005). With respect to wildlife habitat, treatments that are primarily designed to reduce fuels tend to simplify and homogenize the landscape (North et al. 2009). The removal of overstory and understory trees and more uniform spacing of remaining trees will likely result in significant changes in wildlife populations, particularly species that have narrow habitat tolerances in association with habitat features such as canopy cover, understory structure, herbaceous plants (e.g., seed eaters and understory foraging species), fungi, snags and downed logs (Manley 2009). Substantial manipulation of vegetation will likely result in a myriad of short and long-term effects on the composition and structure of plants and animal communities and populations (George and Zack 2001, Bigelow and Manley 2009, Manley 2009). Previous studies in WUI zones in the Basin have found that species responses vary within and among taxonomic groups (Manley et al. 2006, Schlesinger et al. 2008, Sanford et al. 2009). Some species appear to be sensitive to site disturbance (e.g., presence of people and equipment), whereas others are highly sensitive to vegetation structure, and still others are well adapted to a wide variety of environmental conditions. Short-term effects on animals are difficult to predict because the response of species to the disruption will vary depending on the timing and intensity of activities and the sensitivity of each species. Longer-term changes in animal communities and populations resulting from treatments depend heavily on patterns of vegetation recovery following treatment. 5 2.2 PROJECT OBJECTIVES With this study, we aimed to understand the changes to plant and animal communities in response to conventional fuels reductions treatments in the LTBMU. Specifically, our objectives were to: • • • Describe changes in forest structure following hand and mechanical treatments Predict changes in fire behavior following fuel treatments Understand how fuels treatments may change community composition and abundance of small mammals, birds, and ants Understanding how these elements respond to treatments is essential for addressing key management questions identified in the Sierra Nevada Forest Plan Amendment (SNFPA). The data for this project has been collected across a range of treatment types and intensities, in both west shore mixed conifer and east shore Jeffrey pine/red fir forests. The resulting dataset represents the first comprehensive monitoring effort in the Lake Tahoe Basin specifically designed to examine changes in plant and animal communities and fuel loading in response to fuel management activities. 3.0 METHODS 3.1 STUDY SITES We collected data for this project at three general locations in the Lake Tahoe Basin (Figure 1): two on the west shore and one on the east shore of Lake Tahoe. Climate in the Basin is Mediterranean with a summer drought period. The majority of precipitation falls as snow in the winter from December to March and less than 3% falls as rain between May and October. The western portion of the Basin receives greater precipitation and has a higher water balance than the northern and eastern portions of the Basin, resulting in higher plant and animal species richness on the west shore. Mean annual precipitation on the west shore at Tahoe City, CA is 32 in (80 cm) and mean annual snowfall is 190 in (483 cm). Mean annual precipitation on the east shore at Glenbrook, NV is 18 in (46 cm) and mean annual snowfall is 93 in (236 cm). At lake level near Tahoe City, average January high temperature is 42 °F (6 °C). Summers are mild with an average high temperature of 79 °F (26 °C) in August (Western Regional Climate Center 2012). 6 Table 1. Location and characteristics of treatment (T) and control (C) sample units. Treatment units are further identified as Mechanical (M) or Hand (H) thin units. Unit West Shore: BLK 1-4 MCK 13-1 MCK 13-3 TWC 3 WRD 20-16 WRD 20-9 Treatment/ Control Vegetation type HUC 7 watershed T (M) C T (H) C T (M) C T (M) C T (H) C T (M) C Mixed conifer Mixed conifer Mixed conifer Mixed conifer Mixed conifer Mixed conifer Mixed conifer Mixed conifer Mixed conifer Mixed conifer Mixed conifer Mixed conifer Blackwood Creek T (H) C T (H) C Jeffrey pine/red fir Jeffrey pine/red fir Jeffrey pine/red fir Jeffrey pine/red fir Eagle Rock-MaddenHomewood-Quail Eagle Rock-MaddenHomewood-Quail Lower Truckee River Ward Creek Frontal Ward Creek Frontal Slope (%) Elevation (ft) Aspect 17 21 11 10 26 24 6 11 11 11 5 5 6,479 6,542 6,827 6,471 6,609 6,350 6,871 6,916 7,048 7,115 6,972 7,008 S E NW E SE E S S flat flat NW flat 35 42 21 14 7,759 7,644 7,635 7,585 W W S W East shore: REDS WLD North Canyon North Canyon 7 Figure 1. Overview of Upland Fuels sample unit locations in the Lake Tahoe Basin. West Shore Study Sites The six study sites on the western side of the Lake Tahoe Basin were located on United States Forest Service land near Tahoma, California in the lower montane zone between 6,250 and 7,200 ft (1900-2200 m) elevation (Table 1). The lower montane zone in the west Basin consists primarily of mixed conifer forests, comprised of up to six dominant canopy species: red (Abies magnifica var. magnifica) and white fir (A. concolor), Jeffrey (Pinus jeffreyii), lodgepole (P. contorta), and sugar pine (P. lambertiana), and incense cedar (Calocedrus decurrens). Mixed conifer forests on the west shore of the LTBMU occupy approximately 12,300 acres (Murphy and Knopp 2000). Plots were located in four watersheds: Twin Crags (TWC), Ward Creek (WRD), Blackwood Creek (BLK), and McKinney Creek (MCK) (Figures 2 and 3). The six treatment units cover approximately 800 acres in mixed conifer forests. Slopes across the study range from 0 to 40% (Table 1). The Northern Unit Overview (Figure 2) shows more detailed sample plot and control plot locations for the units in TWC, WRD, and BLK. The Southern Overview (Figure 3) shows more detailed sample plot and control plot locations for the units in MCK. 8 Figure 2. Upland Fuels sample unit locations on the northwest shore of Lake Tahoe at Twin Crags (TWC), Ward Creek (WRD), and Blackwood Creek (BLK). 9 Figure 3. Upland Fuels sample plot locations on the southwest shore of Lake Tahoe at McKinney Creek (MCK). East Shore Study Sites The two study sites on the eastern side of the Basin were located on Nevada Division of State Lands (NDSL) property in the North Canyon watershed (Figure 4). Site elevations were all near 7,500 ft and slopes ranged from 14% to greater than 40% (Table 1). The east shore montane zone consists of canopy dominants Jeffrey pine, red fir, and white fir. We chose control sites that were in close proximity to the treatment sites and thereby similar in elevation, forest type and structure. 10 Figure 4. Upland Fuels sample unit locations on the east shore of Lake Tahoe on Nevada Division of State Lands (NDSL) property. 3.2 FUELS REDUCTION TREATMENTS The west shore treatment units were covered under three different NEPA review processes; the Twin Crags units were covered under the North Shore EIS project (1996); the Ward units were part of the Ward Management Area Fuel Hazard Reduction Project Environmental Assessment (2002); and the McKinney and Blackwood units were part of the Quail Vegetation and Fuel Treatment Environmental Assessment (2005). As such, the desired conditions and exact fuels treatment prescriptions varied. For instance, the maximum allowable harvestable tree size increased from 24 to 30 in in 2004 in the SNFPA. Throughout the Basin, fuels reduction treatment prescriptions are applied to retain highest priority conifer species as follows: (1) sugar pine (2) Jeffrey pine (3) white/red fir, incense cedar (4) lodgepole pine. In addition to live tree retention guidelines, all treatment prescriptions specified retention of at least 3 snags and 3 down logs per acre (7.5/ha), both in the largest diameter size class. Our study included four mechanical treatment units, three hand treatment units, and one combined hand/mechanical unit. The hand/mechanical treatment unit (WLD) was ultimately classified as a hand treatment unit for statistical analysis, as the treatments as implemented were most similar in treatment intensity to other hand treatment units. Further details of treatment prescriptions are outlined in Table 2. 11 Three of the treatment units were located in protected activity centers (PACs) for California Spotted Owl and Northern Goshawk. Treatment prescriptions for PACs within the west shore units were prepared in consultation with wildlife biologists in order to maintain or enhance habitat conditions while meeting the objectives of the proposed fuel reduction. The general prescription for mechanical treatment within a unit containing a PAC allows for the retention of larger trees, twice as many snags, two canopy layers and a higher minimum percentage of residual canopy closure. Table 2. General fuels reduction treatment prescriptions by unit. Unit Ownership Treatment Prescription WRD 20-16 USFS Hand thin to 10", pile/burn, PAC MCK 13-1 USFS Hand thin to 14", pile/burn MCK 13-3 USFS Mechanical thin to 30”, masticate and surface spread slash WRD 20-9 USFS Mechanical thin to 24", masticate and surface spread slash BLK 1-4 USFS Mechanical thin to 24", masticate and surface spread slash, PAC TWC 3 USFS Mechanical thin to 24", masticate and surface spread slash, PAC REDS NDSL Hand thin to 24”, pile/burn WLD NDSL Combination hand/mechanical thin to 24”, pile/burn 3.3 SAMPLING DESIGN AND TIMELINE An integrated sampling design was used to collect data on vegetation structure and fuel loads, birds, small mammals and invertebrates. A macroplot of 150 x 330 m was established in a relatively homogeneous and representative portion of each unit. The macroplot provided the grid for 72 small mammal trapping stations, 4 bird survey points, 3 ant pitfall traps, and 8-10 randomly selected vegetation plots (Figure 5). We sampled vegetation in 1 year pre-treatment and 1 year post-treatment, and birds, small mammals, and invertebrates 1-2 years pre-treatment and 2 years post-treatment to examine the impact of fuels treatments on wildlife at these sites (Table 3). 12 Vegetation plots = Bird point counts = Ant bait stations = 1 12 A 150 m F 30 m 330 m Figure 5. Sampling macroplot layout. Each macroplot occupied 5 ha and consisted of 72 small mammal trapping points (every intersection on the grid), 4 bird survey points, 3 ant points, and 8-10 vegetation plots (location of vegetation plots varied by unit). Table 3. Fuels treatment and wildlife sampling timeline for eight paired study sites in the Lake Tahoe Basin. Unit BLK 1-4 Pre 1 2006 MCK13-1 2006 MCK 13-3 2006 RED 2007 TWC3 2006 WLD 2007 WRD 20-16 2006 WRD 20-9 2006 Pre 2 2008 2008 Treatment year 2006 Post 1 2007 Post 2 2008 2006 2007 2008 2009 2010 2011 2007 2008 2009 2008 2009 2010 2008 2009 2010 2007 2007 2008 2009 2007 2008 2009 2010 13 3.4 VEGETATION AND FUELS Vegetation sampling focused on forest structure, understory composition, and fuel loads. Vegetation plot lay-out and sampling protocols were developed in collaboration with the USFS Adaptive Management Services Enterprise Team (AMSET, located on the Tahoe National Forest) and the LTBMU and were based primarily on FIREMON, the Fire Effects Monitoring and Inventory System (Lutes 2006). FIREMON is an Access database software tool that is specifically designed to facilitate analysis of changes in plant communities and fuel loadings in response to management activities over time. The resulting sampling protocols have been used by AMSET to monitor other fuels treatment sites elsewhere in the Basin. Thus, the protocols used in this project are accepted by management agencies in the Basin and will become the primary system to be used for fuels treatment monitoring and associated data management across multiple agencies in the Basin (see www.fire.org). Vegetation sampling plots were fixed-radius circular areas with a 17.84 m (58-ft) radius encompassing 0.1 ha (0.25 ac). Every tree and snag ≥ 6 in (15 cm) DBH was tagged with a unique number and the following information was recorded: species, DBH, total height, decay class (for snags), and live crown ratio. Canopy cover was measured using a GRS site-tube densitometer. Surface and ground fuels were sampled on four, 20-m transects in each inventory plot using the line-intercept method (Brown and Johnston 1976). Herb and shrub percent cover were measured in five 0.25-m2 quadrats along each transect. Detailed data collection parameters for vegetation and fuels can be found in Appendix A. 3.5 WILDLIFE We collected data on both vertebrate and invertebrate response to fuels treatments. Birds and small mammals were the focus of vertebrate sampling because they have complementary sensitivities to the effects of fuels treatments. They also serve as the primary prey for upper trophic level species of special status in the Basin, namely the California Spotted Owl, Northern Goshawk, and American marten (Martes americana). Small mammals are closely tied to specific features of forest vegetation structure and they are primarily year-round residents. Birds are more mobile, but most species have relatively narrow environmental conditions in which they can successfully breed, and they are also dependent upon both overstory and understory conditions. Ants provide a variety of ecological benefits, including soil aeration, nutrient cycling, and enhanced water infiltration. Thus, the combined responses of plants, vertebrates, and invertebrates provide rich information about ecological conditions. We trapped small mammals to determine species occupancy and abundance. We sampled small mammals by placing one Tomahawk™ live trap (12.5 x 12.5 x 40 cm) and one extra large Sherman live trap (10 x 11.5 x 38 cm) at each of 72 trap stations (6 x 12 stations, 30 m apart = 150 x 330 m grid = 5 ha area) (Figure 5). We attached Tomahawk™ traps to the trunk of trees > 20 in (50 cm) DBH, 1.5-2.0 m above ground. We selected trees that were as close to the trap station as possible, ideally within 5 m. We covered the traps with a tarp around the outside and placed a nest box (10 x 10 x 6 cm cardboard) at the back of the trap with polystyrene for warmth. We placed Sherman traps on the ground at the base of trees or along larger logs or under shrubs. Traps were securely placed such that they did not rock or move when an animal entered. We covered each Sherman trap with natural materials to insulate traps from the sun and rain, and placed polystyrene in the back of the trap for warmth. We used a bait mixture of oats, bird seed and raisins. We attempted to include peanut butter and molasses, following the general formula used by Carey et al. (1991), but excessive bear damage required that these ingredients were eliminated or 14 reduced. Traps were set, baited, and locked open for a minimum of three nights before trapping began, then opened for five days, starting with traps being set in the late afternoon/early evening prior to the first trap evening (just before dusk). We checked traps twice per day, generally before 10 a.m. for morning checks and after 4 p.m. for afternoon checks. All traps were removed the morning after the fourth night. We calculated trapping effort by correcting for traps rendered unavailable for some (0.5; disturbed, robbed, or sprung) or all (1.0; missing or destroyed) of a trapping occasion. We identified all individuals captured to species and marked each animal using uniquely numbered ear tags and recorded data on sex, age (juvenile or adult), and weight. Ear, leg, or tail measurements were taken on individuals whose identification was in question. Bird point count stations were located at four points along the center line of the sample plot (Figure 5). We conducted three point count visits to each sample plot between late May and early July. We used a 10 minute survey period and recorded the distance to all birds using 20 m increments. Paired sites were sampled within two days of one another, and multiple observers were rotated among visits across sites in order to minimize observer bias. Surveys began at least 15 minutes after sunrise and were completed before 9:30 a.m. We avoided conducting surveys during conditions which limited bird activity or detectability (e.g., strong wind or rain). Composition and relative abundance of ants were characterized with a grid of pitfall traps at 40 sites. We arranged 12 traps in a 40 x 40 m grid, with four traps, spaced 10m apart, along each of three 40 m transects that were oriented north-south and centered on the center point of each plot; the three transects were separated by 20 m (Bestelmeyer and Wiens 1996, Andersen 1997). Pitfall traps consisted of 6.5 cm diameter (120 ml) plastic cups with approximately 25 ml of propylene glycol. Traps were left open for seven days and then the contents were collected. Ants were identified to species whenever possible, and abundance of each species and number of species were summed over all transects to obtain total abundance and species richness for each site. 3.6 DATA ANALYSIS 3.6.1 Vegetation and Fuels We characterized forest conditions before and after treatment by calculating values for the following variables: quadratic mean tree diameter (hereafter QMD), percent canopy cover, tree density, tree basal area, snag density, and the amount of coarse woody debris (hereafter CWD). We calculated tree density by species and diameter class to assess species composition and vegetation structure. The composition and cover of herbaceous and shrub species were also characterized and included in estimates of fuels. Understory vegetation is a critical component of wildlife habitat and it can strongly influence fire behavior. We analyzed changes in these vegetation variables using PROC MIXED in SAS 9.2 (SAS Institute 2008). As we were not interested in main effects (treatment or timing) in isolation, but rather the interaction of main effects, we included the interaction of treatment and timing as our only fixed effect. Treatment unit was included as a random effect, and sampling plot was nested within unit. To examine differences between treatment types, we included an ESTIMATE statement. This statement provides estimated differences in response variables between specified treatment types and time periods. 15 Based on the tree measurements from each plot we developed pre- and post-treatment tree lists for each plot. These tree lists, along with surface fuel information were entered into Fuels Management Analyst Plus (FMA) to simulate fire behavior and predict fire-caused tree mortality. FMA uses published equations to model fire behavior and effects based on user-input weather and fuel moisture conditions (Carlton 2005, Stephens and Moghaddas 2005, Stephens et al. 2009). Often the weather and fuel moisture conditions are based on higher percentile weather (e.g., 95th or 97th), however in our simulations we chose to use the observed conditions from a recent “problem” fire that occurred in the Lake Tahoe Basin in 2007 (Angora fire), as these are the conditions that are known to have resulted in exacerbated fire behavior and effects. The actual weather values used were as follows: fuel moistures 1, 2, and 5% for 1hr, 10-hr, and 100-hr timelag classes, respectively; wind speed (6.3m) 33 km hr-1; and air temperature 26.6˚C. For surface fuels, which are represented in the fire behavior simulations as surface fuel models, we attempted to bracket the uncertainty associated with assigning surface fuel models (Collins et al. 2011) by running fire behavior simulations under two fuel model scenarios, a higher and lower option, for each plot pre- and post-treatment. Fuel model assignments were made at the treatment unit level and were based on several factors. Pre-treatment fuel model assignments were based on unit-level average fuel loads and fuel depths. These varied slightly among treatment units, and as a result pre-treatment fuel models were fairly similar across units (Table 5). Post-treatment fuel model assignments were also based on average fuel loads and depths; however, for masticated stands, which had greater fuel depths (Table 5), assignments followed the recommendations in Knapp et al. (2011). According to Knapp et al. (2011), the two fuel models we used (NFFL 9 and SB 202) bracketed observed fire behavior fairly well, balancing over/under prediction of rate of spread vs. flame length. We tested for both treatment and treatment intensity effects (basal area change from pre- to posttreatment) on predicted fire-caused mortality using a mixed model analysis (PROC MIXED; SAS Institute 2008). We used two variables to capture predicted mortality for pre- and post-treatment stand conditions: fire-killed basal area proportion and number of fire-killed large trees (> 61.0 cm DBH). Treatment unit was considered a random effect in the model. We investigated diagnostic plots of the residuals for compliance with normality and homogeneity of variance assumptions. Fire-killed basal area proportion met these assumptions, while number of fire-killed large trees had to be square roottransformed to meet assumptions. 3.6.2 Wildlife We described species abundance, richness, and composition for all small mammal and bird species detected on our surveys. To examine small mammal response to fuel treatment, we examined communitylevel changes in species richness and composition. In addition, we looked at changes in species abundance following treatment implementation. Because of difficulties with the identification of individual species of chipmunks (Tamias sp.), we also pooled abundance data for all chipmunk species to increase sample size and examine genus-level response to treatment (Converse et al. 2006, Bagne and Finch 2010). Abundance was calculated using the number of unique individuals captured per 100 trap nights. We analyzed species- and genus-specific responses to treatment using PROC MIXED in SAS 9.2 (SAS Institute 2008). In our base model, we included abundance as the response variable, and the interaction of treatment and timing as our fixed effect. Treatment unit was included as a random effect. Furthermore, because wildlife populations, and particularly small mammal populations, show strong annual variability, and because treatments were implemented in different years, we ran an additional 16 model which included year as a random effect. We compared model fit between the base and sampling year models using AICc scores (Burnham and Anderson 2002), and report here the results from the bestsupported model (lowest AICc value). Additionally, we ran several additional models which grouped treatment types (i.e., both hand and mechanical thins lumped under a single “treatment” category) and time since treatment (both post-treatment years combined); however, all models which simplified treatment type or timing were poorly supported, for all species considered. We excluded any species that were captured less than once per every 1000 trap nights (35 captures total) over the course of our study. We calculated species richness by tallying all the unique species captured at a site over the course of the trapping period. Bird species richness and abundance metrics were calculated for each site. Bird species richness was based on all species encountered over the course of all three visits to a site during a year. We calculated bird abundance as the mean number of individuals detected across all four count stations and three visits (12 counts total) for each site and year. We included detections within 60m of the point count station, in order to minimize double-counting of individuals and obtain more reliable estimates of abundance. We analyzed species-specific response to treatment as outlined above for small mammals. We did not analyze response for any species that were detected fewer than 10 times. We employed several different methods to measure community similarity, diversity, and evenness. For these calculations, we pooled the data from all 8 treatment sites. For bird species, we used the average abundance recorded within a year averaged across pre- and post- treatment years. For small mammal species, we used the total number of uniquely marked adult individuals captured during a five-night trapping session and averaged across pre- and post-treatment periods. In order to measure and compare the degree of association or similarity in bird and small mammal community composition at sites before and after treatments were implemented, we used the Renkonen index (P). Also known as the Percentage Similarity index, this is a quantitative index that accounts for differences in the abundance of species (Wolda 1981). This index ranges from 0 to 1, with 0 indicating no overlap, and 1 indicating complete similarity. We used Shannon’s diversity index (Shannon 1949) to examine changes in both species richness and dominance at the treatment sites before and after treatment. We also separately examined community evenness before and after treatment using Shannon’s Evenness index (Pielou 1966). In determining the effect of treatment on ants, we used presence/absence data, as opposed to abundance data, because ant abundance is sensitive to distance to nest site. A species was considered present at a site if one or more individuals were detected on the grid. For sites that were sampled more than one time before and after treatment, we selected the site that was sampled in the time interval closest to the implementation of treatment. We compared the number of sites with each species in each treatment category (control, hand and mechanical) before and after treatment with Fisher’s Exact Test. We also compared species richness of ants with multi-factor, repeated measures ANOVA to see if sites differed in richness before and after treatment or by treatment type. We looked at the interaction of timing (pre- or post-treatment) and treatment type to compare control sites to treated sites before and after implementation. We also included year in our analysis to determine if changes were due to treatment effects or year effects. Finally, we examined changes in ant community diversity and evenness using the methods outlined above for vertebrates. 17 4.0 RESULTS 4.1 VEGETATION Pre-treatment forests at our study sites were largely dominated by white and red fir, with one of our eastshore sites (WLD) dominated primarily by pine (Figure 6). In addition, pre-treatment forests were composed of a roughly even mix of pole-sized (6-12” dbh) and small (12-24”) trees (Figure 7). While treatment prescription and intensity varied by site, treatments generally targeted smaller-diameter trees, and preferentially removed firs given their fire susceptibility and ability to serve as ladder fuels. Although still fir-dominated, post-treatment forests showed a greater composition of pines (Figure 6), and were generally dominated by 12-24” trees (Figure 7). While the quantity of medium and large (>24”) trees was little changed, they became a more prominent component of post-treatment forests, given the preferential removal of trees in the smaller size classes. 600 Pre-treatment Post-Treatment 500 Trees/ha 400 300 200 100 MCK 13-1 RED WLD WRD 20-16 Hand BLK 1-4 MCK 13-3 TWC 3 Cedar Pine Fir Cedar Pine Fir Cedar Pine Fir Cedar Pine Fir Cedar Pine Fir Cedar Pine Fir Cedar Pine Fir Cedar Pine Fir 0 WRD 20-9 Mechanical Figure 6. Pre- and post-treatment mean tree density for groups of like trees: fir, cedar, and pine at eight treatment units in the Lake Tahoe Basin. 18 350 Pre-treatment 300 Post-Treatment Trees/ha 250 200 150 100 50 MCK 13-1 RED WLD WRD 20-16 Hand BLK 1-4 MCK 13-3 TWC 3 > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" 0 WRD 20-9 Mechanical Figure 7. Pre- and post-treatment mean tree density by diameter class in eight treatment units in the Lake Tahoe Basin. Mechanical fuels treatments led to substantial reduction in basal area, while hand-thin units showed little change in basal area (Figure 8). The reduction in basal area was significant on mechanically treated plots (F2,144 = 46.88, p < 0.001; Table 4). Post-treatment basal area showed wide variation among plots, ranging from 123.3 to 330.2 ft2/ac, with an average of 205.8 ft2/ac. The preferential removal of smaller-diameter trees resulted in an increase in average tree diameter (as measured by QMD) on all plots (Figure 9). The increase in QMD varied substantially between plots, however, ranging from as little as 0.17” to greater than 5” on several plots. QMD was significantly affected by treatment (F2,137 = 119.70, p < 0.01). Both treatment types significantly increased QMD, and mechanical treatments resulted in a significantly greater increase than did hand treatments (Table 4). 19 Basal Area (ft2/ac) 500 450 400 350 300 250 200 150 100 50 0 Pre-treatment Post-treatment Control MCK 13-1 RED WLD WRD 20-16 BLK 1-4 Hand MCK 13-3 TWC 3 WRD 20-9 Mechanical Figure 8. Pre- and post-treatment basal area on eight treatment units and corresponding control units in the Lake Tahoe Basin. Quadratic Mean Diameter (in.) 25 Pre-treatment Post-treatment 20 Control 15 10 5 0 MCK 13-1 RED WLD WRD 20-16 Hand BLK 1-4 MCK 13-3 TWC 3 WRD 20-9 Mechanical Figure 9. Pre- and post-treatment quadratic mean diameter of trees in eight treatment units and corresponding control units in the Lake Tahoe Basin. Snags showed a similar pattern to that of live trees following treatment. Prior to fuels reduction, polesized and small snags were roughly even in abundance on most plots (Figure 10). Because of their associated fire risk and lower value to wildlife (Dodson and Peterson 2009), smaller diameter snags were removed most frequently, and 12-24” snags were most common in post-treatment forests. Snags > 24” also appeared to be reduced on most plots, though this may have been the result of wind or natural degradation, rather than active removal of larger snags. 20 250 Pre-treatment Post-Treatment Snags/ha 200 150 100 50 MCK 13-1 RED WLD WRD 20-16 BLK 1-4 Hand MCK 13-3 TWC 3 > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" > 24" 12-24" 6-12" 0 WRD 20-9 Mechanical Figure 10. Pre- and post-treatment mean density by diameter class of snags in eight treatment units in the Lake Tahoe Basin. % Canopy Cover Given the removal of both live trees and snags, there was a predictable decrease in canopy cover at all treatment sites (Figure 11; F2,149 = 20.89, p < 0.01). However, there was considerable variation in this measure, with treatments resulting in between 4 and 22.8% reduction in canopy cover, depending on the treatment unit. Post-treatment canopy coverage ranged from 31.6 to 52%, with a mean of 41.2%. This reduction was significant for both hand and mechanical treatments (Table 4). 80 Pre-treatment 70 Post-treatment 60 Control 50 40 30 20 10 0 MCK 13-1 RED WLD WRD 20-16 Hand BLK 1-4 MCK 13-3 TWC 3 WRD 20-9 Mechanical Figure 11. Pre- and post-treatment average canopy cover on eight treatment units and corresponding control units in the Lake Tahoe Basin. 21 Understory vegetation was also reduced in extent following treatment at all sites. Average shrub cover was reduced from 16.9% to 9.4% following treatment (Figure 12; F2,123 = 6.81, p = 0.002). This reduction in shrub cover was significant for only mechanical treatments, while hand-thin units experienced a moderate, non-significant decline in shrub cover (Table 4). Herbaceous cover was limited prior to treatment, averaging 2.8% across all treatment sites, and was reduced following treatment to 1.2% (Figure 13; F2,123 = 0.09, p = 0.91), though this change was not significant for either treatment type (Table 4). Another important component of forest understory structure, coarse woody debris, was significantly reduced in volume following both treatment types (Figure 14; F2,147 = 6.50, p = 0.002). Mechanical treatments removed a greater volume of CWD than did hand treatments (Table 4), likely a result of mastication of residual materials. 35 % Shrub Cover Pre-treatment 30 Post-treatment 25 Control 20 15 10 5 0 MCK 13-1 RED WLD WRD 20-16 Hand BLK 1-4 MCK 13-3 TWC 3 WRD 20-9 Mechanical Figure 12. Average shrub cover on eight treatment units and corresponding control units before and after treatments in the Lake Tahoe Basin. 22 % Herbaceous Cover 14 Pre-treatment 12 Post-treatment 10 Control 8 6 4 2 0 MCK 13-1 RED WLD WRD 20-16 BLK 1-4 Hand MCK 13-3 TWC 3 WRD 20-9 Mechanical Coarse Woody Debris (m3/ha) Figure 13. Average herbaceous cover on eight treatment units and corresponding control units before and after treatments in the Lake Tahoe Basin. 1200 Pre-treatment 1000 Post-treatment Control 800 600 400 200 0 MCK 13-1 RED WLD WRD 20-16 Hand BLK 1-4 MCK 13-3 TWC 3 WRD 20-9 Mechanical Figure 14. Average coarse woody debris (CWD) density, in m3/ha, on eight treatment units and corresponding control units before and after treatment in the Lake Tahoe Basin. 23 Table 4. Relative changes in vegetation variables on mechanical and hand units, relative to control units, as a result of fuels reduction treatments. Mechanical Measurement Basal Area (ft2/ac) Estimate t Hand p Estimate t p -118.91 ± 12.56 -9.47 < 0.001 -15.58 ± 13.27 -1.17 0.24 4.61 ± 0.31 14.82 < 0.001 2.99 ± 0.35 8.60 < 0.001 -16.32 ± 2.61 -6.26 < 0.001 -9.75 ± 2.70 -3.61 < 0.001 Shrub Cover (%) -7.05 ± 1.91 -3.69 < 0.001 -2.64 ± 1.98 -1.33 0.18 Herbaceous Cover (%) -0.32 ± 0.75 -0.42 0.67 -0.09 ± 0.72 -0.12 0.90 -270.22 ± 81.91 -3.30 0.001 -211.97 ± 86.52 -2.45 0.02 QMD (in.) Canopy Cover (%) 3 CWD (m /ha) 4.2 FUELS There were inconsistent responses to treatment in fine woody surface fuel loads and fuel depths across treatment units (Table 5). In 5 of the 8 units there was nearly no change in fine woody fuel loads from pre- to post-treatment (Table 5). The one consistent trend was a noticeable increase in fuel depth for all masticated units from pre- to post-treatment. Fuel depth in non-masticated units was relatively unchanged (Table 5). 24 Table 5. Treatment unit average fine woody surface fuel loads (0-7.62 cm diameter or 1-100h) and fuel depths based on plot measurements. Chosen surface fuel models for low and high fire behavior conditions are also reported. We used fuel models from both the original set of 13 described by Anderson (1982) and expanded set of 40 described by Scott and Burgan (2005). Surface fuel model Fine woody surface Fuel depth (cm) Number -1 fuel loads (Mg ha ) Pre Post Unit of plots Pre Post Pre Post Low High Low High BLK 1-4* 10 7.1 8.0 3.1 5.4 201 10 9 202 WRD 20-9* 10 8.4 8.2 2.3 5.6 201 10 9 202 MCK 13-3* 10 10.9 7.4 3.1 5.8 10 165 9 202 TWC 3* 10 6.5 7.5 1.1 5.2 201 10 9 202 MCK 13-1 9 7.7 10.8 2.9 2.4 201 10 201 165 WRD 20-16 10 6.5 7.6 2.2 2.0 201 10 201 10 RED 8 6.3 6.8 2.0 3.0 201 10 201 10 WLD 8 8.7 6.0 3.2 4.7 201 165 201 10 *masticated units with post-treatment fuel models based on recommendations in Knapp et al. (2011) Treatment intensity, defined as the average basal area change from pre- to post-treatment for all plots in a treatment unit, varied from 0 to 25 m2/ha across the 8 treatment units (Figure 15). Treatment units fell in three broad categories with respect to treatment intensity: 1) BLK 1-4, WRD 20-9, and MCK13-3 had the highest treatment intensity (all > 20 m2ha-1 change), TWC 3 and MCK 13-1 had intermediate treatment intensity (~11 m2ha-1 change), while WRD 20-16, WLD, and RED had the lowest intensity (0-2 m2ha-1 change). The highest intensity treatments noticeably reduced tree density across all but the largest tree size classes (> 61.0 cm DBH) (Figure 16). The moderate intensity treatments slightly reduced tree density in the 30.5 to 61.0 cm DBH class, but considerably reduced density in the two smaller size classes (2-15.1 and 15.2-30.4 cm DBH). Tree density reductions in the lowest intensity treatments were primarily in the two smaller size classes (Figure 16). Canopy cover and canopy bulk density from pre- to post-treatment followed a similar trend, i.e., greater reductions in units with higher treatment intensities, with a couple notable exceptions. In TWC 3 and WRD 20-16 canopy cover was relatively similar from pre- to post-treatment (Figure 17), despite losing over half of the trees in the 15.2 to 30.4 cm DBH class (Figure 16). Canopy base height increased after treatment in all treatment units but one, WLD (Figure 17). The largest increases in canopy base height were in the three units with the highest treatment intensity (Figure 17). The mixed model analysis indicated a strong treatment effect (P<0.001) for both response variables: firekilled basal area proportion and number of fire-killed large trees (> 61.0 cm DBH). For variables there was a considerable decrease in predicted mortality following treatment (Figures 15 and 16). There was no evidence for a treatment intensity effect on fire-killed basal area proportion (P=0.27) and marginal evidence for a treatment intensity effect on the number of fire-killed large trees (P=0.09), indicating more intense treatments result in slightly less large trees being fire-killed. The interaction between treatment and treatment intensity was only significant for the model explaining fire-killed basal area proportion (P<0.01). 25 Basal area (m2/ha) 70 live 60 fire-killed 50 40 30 20 10 0 BLK 1-4 WRD 20-9 MCK 13-3 TWC 3 MCK 13-1 WRD 20-16 WLD RED Decreasing Treatment Intensity → Figure 15. Average basal area for each treatment unit pre- and post-treatment (post indicated by horizontal line fill). Predicted mortality under modeled wildfire conditions (see Methods) is shown within the bars for each unit. 26 2.5 to 15.1 cm 30.5 to 61.0 cm 1000 300 live 250 800 fire-killed 200 600 150 400 Trees/ha 100 200 50 0 0 BLK 1- WRD MCK TWC 3 MCK WRD WLD RED 4 20-9 13-3 13-1 20-16 BLK 1- WRD MCK TWC 3 MCK WRD WLD RED 4 20-9 13-3 13-1 20-16 15.1 to 30.5 cm > 61.0 cm 300 60 250 50 200 40 150 30 100 20 50 10 0 0 BLK 1- WRD MCK TWC 3 MCK WRD WLD RED 4 20-9 13-3 13-1 20-16 BLK 1- WRD MCK TWC 3 MCK WRD WLD RED 4 20-9 13-3 13-1 20-16 Decreasing Treatment Intensity → Figure 16. Average tree density by diameter-at-breast height class for each treatment unit pre- and posttreatment (post indicated by horizontal line fill). Predicted mortality under modeled wildfire conditions (see Methods) is shown within the bars for each unit. 27 % Cover Canopy Cover 70 60 50 40 30 20 10 0 BLK 1-4 WRD 20-9 MCK 13-3 TWC 3 MCK 13-1 WRD 20-16 WLD RED WLD RED WLD RED Canopy Bulk Density 0.10 kg/m3 0.08 0.06 0.04 0.02 0.00 BLK 1-4 WRD 20-9 MCK 13-3 TWC 3 MCK 13-1 WRD 20-16 Canopy Base Height 10 Meters 8 6 4 2 0 BLK 1-4 WRD 20-9 MCK 13-3 TWC 3 MCK 13-1 WRD 20-16 Decreasing Treatment Intensity → Figure 17. Average canopy characteristics for each treatment unit pre- and post-treatment (post indicated by horizontal line fill). 28 4.3 SMALL MAMMALS We captured 10682 small mammals from 16 species during the course of our project (Appendix B). At the community level, there was minimal change in response to treatment. Treatment units had 14 species in both pre- and post-treatment samples. Two species present in the pre-treatment samples (pinyon mouse, Peromyscus truei, and brush mouse, Peromyscus boylii) were not detected during post-treatment sampling. Likewise, there were two species detected during post-treatment sampling that were not detected prior to treatment (least chipmunk, Tamias minimus, and bushy-tailed woodrat, Neotoma cinerea). However, the identity of the least chipmunk should be confirmed with genetic data given difficulties in identification and location of the Tahoe Basin on the outer extent of the species range. The Renkonen index indicated 82.4% similarity between pre- and post-treatment samples across treatment units, when accounting for changes in both species composition and abundance. Shannon’s Diversity index remained virtually unchanged, with a very minor increase following treatment (Table 6). Likewise, there was only a small change in community evenness following treatment, as indicated by Shannon’s Evenness Index suggesting that the proportion of the community represented by different species remained similar. When considered as a whole, the small mammal community showed an increase in abundance in response to both hand and mechanical treatments, in both sampling years following treatment. Abundance increased between 12% and 28%, depending on year and treatment type, though these changes were nonsignificant (all p > 0.1). At the species level, we had sufficient data from 10 species to analyze their response to treatment (Table 7). Although several species exhibited substantial changes in abundance following treatment, these changes were statistically significant for only 2 species. In the first year post-treatment, deer mice (Peromyscus maniculatus) declined significantly on mechanical thin units, and lodgepole chipmunks (Tamias speciosus) declined significantly on hand-thinned units. Although all other species-level results were non-significant, there were several consistent patterns observed. In particular, yellow-pine chipmunks (Tamias amoenus) increased in abundance on both mechanical and hand treatment units during both years of post-treatment surveys, with increases ranging from 15 to 130%. Additionally, northern flying squirrels (Glaucomys sabrinus) declined on all treatment units during both years of the study, with declines ranging from 33 to 70%. Finally, because of challenges with chipmunk identification, we pooled data from all chipmunk species to examine the group’s response to treatment. When grouped, chipmunks increased in abundance in both years following treatment on both hand and mechanical units, with increases between 28.3 and 61.3%. However, the only statistically significant increase was in the 2nd year post-treatment on mechanical sites. 29 Small mammals/100 trap nights 40 Year 1 30 Year 2 20 10 0 -10 -20 -30 -40 -50 C T C MCK 13-1 T C TWC 3 T C T C WRD 20-16 WRD 20-9 T BLK 1-4 C T C MCK 13-3 Hand T C RED T WLD Mechanical Small mammal species richness Figure 18. Change in mean small mammal abundance in years 1 and 2 post-treatment, relative to pretreatment years, at paired control (C) and treatment (T) sites in the Lake Tahoe Basin. Means are corrected for differences in trapping effort and reported as # captures/100 trap nights. 4 3 2 1 0 -1 -2 -3 -4 -5 Year 1 Year 2 C T MCK 13-1 C T TWC 3 C T WRD 20-16 C T WRD 20-9 Hand C T BLK 1-4 C T MCK 13-3 C T RED C T WLD Mechanical Figure 19. Change in small mammal species richness in years 1 and 2 post-treatment, relative to pretreatment years, at paired control (C) and treatment (T) sites in the Lake Tahoe Basin. 30 Table 6. Calculation of diversity, similarity and evenness using the total number of individual mammals (by species) captured before and after fuels reduction in the Lake Tahoe Basin, 2006-2011. Number of species Number of species in common (j) Renkonen index (P) Shannon diversity (H') Shannon evenness (E) Treatment Pre-treatment Post-treatment 14 14 12 0.824 1.87 1.93 0.71 0.73 Control Pre-treatment Post-treatment 10 15 7 0.308 1.01 1.14 0.44 0.42 Table 7. Percent change in abundance, relative to control sites, following fuels reduction treatments among small mammal species detected in the Lake Tahoe Basin, 2006-2011. Species are listed in order of abundance. Mechanical Hand % Change Year 1 % Change Year 2 Direction of Change % Change Year 1 % Change Year 2 Direction of Change Deer mouse Long-eared chipmunk Allen’s chipmunk Yellow-pine chipmunk Lodgepole chipmunk -67.5 2.9 38.4 113.8 54.8 2.8 19.4 38.1 129.5 -13.6 Mixed Positive Mixed Positive Mixed 4.4 21.5 -19.4 72.2 -260.8 30.0 0.3 -14.4 15.5 36.9 Positive Positive Mixed Positive Mixed Golden-mantled ground squirrel Douglas’ squirrel Northern flying squirrel California ground squirrel Least chipmunk All Chipmunks All Species 11.8 129.5 -33.2 -37.3 -26.4 61.3 12.0 -18.9 -0.4 -70.2 127.0 † 48.4 27.6 Mixed Mixed Negative Negative Mixed Positive Positive -29.9 -8.0 -47.7 -58.9 56.2 28.3 23.6 22.2 -74.3 -47.6 -74.5 † 29.3 15.8 Mixed Negative Negative Mixed Positive Positive Positive Common Name Bold = p < 0.05; † = No detections on control plots, unable to calculate % increase 4.4 BIRDS Across the six years of our study, we observed 8110 individuals from 63 species of birds (Appendix C). Species composition of the bird community showed minimal change following treatment. We detected 53 species on treatment units prior to treatment, and 57 species following treatment; 47 of these species were found both before and after treatment. The Renkonen index, which examines both species composition and relative abundance, indicated 85.6% similarity between pre- and post-treatment bird communities (Table 8), suggesting that these treatments did not produce substantial changes at the community level. Similarly, there were only small changes in both Shannon’s Diversity index (3.27 pre-treatment, 3.28 post-treatment) and Shannon’s Evenness index (0.82 before treatment, 0.81 following treatment), 31 indicating that there were only minimal changes in community diversity and relative abundance following treatment. On control sites, all community metrics were very similar to those found on treatment sites (Table 8), further indicating that treatments produced minimal change to the bird community. We analyzed changes in abundance of 36 bird species for which we had greater than 10 observations (Table 9). In spite of substantial changes in abundance following treatment for many species, few of these differences were statistically significant. The species which showed significant (p < 0.05) positive responses to treatment were Dark-eyed Junco (Junco hyemalis), Dusky Flycatcher (Empidonax oberholseri), Evening Grosbeak (Coccothraustes vespertinus), and Red Crossbill (Loxia curvirostra). Golden-crowned Kinglet (Regulus satrapa), Nashville Warbler (Oreothlypis ruficapilla), and Redbreasted Nuthatch (Sitta canadensis) all had significant negative responses to treatment in one or both years (Table 9). Although few species exhibited statistically significant responses, many species exhibited consistent responses to treatment in both years following treatment. In response to mechanical treatment, 13 species increased both years following treatment, while 12 species declined in both years following treatment. Similarly, with hand treatments, 13 species responded positively in both years post-treatment, while 10 species declined in both years. When all species were grouped for analysis, abundance of the entire bird community declined slightly, but non-significantly (all p > 0.4) on both hand and mechanical treatment plots during both years of posttreatment sampling. Declines ranged from 4.8% to 9.5% relative to control plots, depending on year and treatment type (Table 9). Number of Individuals 8 Year 1 6 Year 2 4 2 0 -2 -4 -6 C T MCK 13-1 C T TWC 3 C T C T C WRD 20-16 WRD 20-9 T BLK 1-4 Hand C T MCK 13-3 C T RED C T WLD Mechanical Figure 20. Change in mean bird abundance in years 1 and 2 post-treatment, relative to pre-treatment years, at paired control (C) and treatment (T) sites in the Lake Tahoe Basin. 32 8 Year 1 6 Bird Species Richness Year 2 4 2 0 -2 -4 -6 -8 C T MCK 13-1 C T TWC 3 C T C T C WRD 20-16 WRD 20-9 T BLK 1-4 C T MCK 13-3 Hand C T C RED T WLD Mechanical Figure 21. Change in bird species richness in years 1 and 2 post-treatment, relative to pre-treatment years, at paired control (C) and treatment (T) sites in the Lake Tahoe Basin. Table 8. Calculation of similarity, diversity, and evenness using total bird detections before and after fuels reduction in the Lake Tahoe Basin, 2006-2011. Number of species Number of species in common (j) Renkonen index (P) Shannon diversity (H') Shannon evenness (E) Treatment Pre-treatment Post-treatment 53 57 47 0.856 3.27 3.28 0.82 0.81 Control Pre-treatment Post-treatment 52 56 42 0.901 3.27 3.26 0.83 0.81 Table 9. Change in abundance of bird species following mechanical- and hand-thin fuels treatments, relative to control sites, in the Lake Tahoe Basin. Results are presented for all species with > 10 detections, and species are listed in order of abundance. Mechanical Common Name Mountain Chickadee Red-breasted Nuthatch Yellow-rumped Warbler Golden-crowned Kinglet Dark-eyed Junco Western Tanager Dusky Flycatcher Hand % Change Year 1 % Change Year 2 Direction of Change % Change Year 1 % Change Year 2 Direction of Change -20.4 -33.4 -8.0 -50.9 87.8 -8.2 -4.7 -27.7 -61.7 14.9 -29.6 74.0 -32.2 21.1 Negative Negative Mixed Negative Positive Negative Mixed 24.7 12.6 14.4 -39.6 -68.8 15.7 21.5 28.9 -14.8 -5.3 -24.1 -36.1 7.1 102.0 Positive Mixed Mixed Negative Negative Positive Positive 33 Table 9 (continued). Mechanical Common Name Evening Grosbeak Steller's Jay Fox Sparrow Nashville Warbler Brown Creeper American Robin Hermit Thrush Warbling Vireo Pine Siskin Brown-headed Cowbird Western Wood-Pewee White-headed Woodpecker Cassin's Finch Townsend's Solitaire Hairy Woodpecker Hermit Warbler MacGillivray's Warbler White-breasted Nuthatch Red Crossbill Cassin's Vireo Clark's Nutcracker Northern Flicker Red-breasted Sapsucker Williamson's Sapsucker Chipping Sparrow Black-backed Woodpecker Band-tailed Pigeon Ruby-crowned Kinglet Pygmy Nuthatch All Species Hand % Change Year 1 % Change Year 2 Direction of Change % Change Year 1 % Change Year 2 Direction of Change 182.7 -5.6 -68.0 -149.9 -2.2 125.9 -90.0 20.6 -79.4 22.1 173.9 -75.3 20.4 123.2 27.6 -51.3 -75.7 45.8 † 13.6 -99.4 -53.0 -315.9 58.6 220.5 94.3 -237.8 35.5 † -9.5 -59.6 -24.2 -88.9 -140.3 18.8 101.5 -88.7 55.5 -53.9 72.3 554.8 20.0 71.4 14.4 32.4 -82.5 12.8 6.9 71.4 226.6 159.7 -81.3 143.2 -172.0 -8.7 105.9 -703.8 † 492.4 -8.6 Mixed Negative Negative Negative Mixed Positive Negative Positive Negative Positive Positive Mixed Positive Positive Positive Negative Mixed Positive Positive Positive Mixed Negative Mixed Mixed Mixed Positive Negative Positive Positive Negative 68.7 -30.6 40.2 -233.7 24.1 -35.9 28.0 36.5 -42.5 -20.9 -9.9 72.2 -6.3 143.9 -22.2 -69.5 -9.3 15.2 † -45.7 202.3 -68.9 -106.9 -5.0 166.6 191.9 6.8 84.1 † -4.8 213.8 -34.3 0.3 -322.8 -12.1 -39.5 -35.2 65.1 -75.6 -42.1 396.2 38.5 27.3 27.1 491.2 28.3 220.0 -109.0 21.9 163.5 -180.8 -87.3 -35.9 -4.0 158.2 497.9 540.0 † 80.5 -7.6 Positive Negative Positive Negative Mixed Negative Mixed Positive Negative Negative Mixed Positive Mixed Positive Mixed Mixed Mixed Mixed Positive Mixed Mixed Negative Negative Negative Positive Positive Positive Positive Positive Negative Bold = p < 0.05; † = No detections on control plots, unable to calculate % increase 4.5 INVERTEBRATES A total of 38 species of ants from 10 genera were detected over the entire study period. Species detected included members of several ecologically important functional groups. These groups included aerators (species that create tunnels that aerate soils and aid in water infiltration), decomposers (ants that build tunnels through woody debris and aid in nutrient release), compilers (species that build thatch mounds that increase soil nutrient availability), and generalists (species that may fill any of the above roles in certain contexts; Sanford et al. 2009). 34 Species composition of the ant community changed following treatment. We detected 22 species on treatment units prior to treatment, and 25 species following treatment; 15 of these species were found both before and after treatment. The Renkonen index, which examines both species composition and relative abundance, indicated 68.6% similarity between pre- and post-treatment ant communities (Table 10), suggesting that these treatments did produce substantial changes at the community level. Similarly, there were changes in both Shannon’s Diversity index (2.34 pre-treatment, 1.79 post-treatment) and Shannon’s Evenness index (0.76 before treatment, 0.56 following treatment), indicating that there were changes in community diversity and relative abundance following treatment. This pattern was also observed when on control sites, though there was greater similarity between pre- and post-treatment communities on the control sites. Table 10. Calculation of similarity, diversity, and evenness using total ant detections before and after fuels reduction in the Lake Tahoe Basin, 2006-2011. Number of species Number of species in common (j) Renkonen index (P) Shannon diversity (H') Shannon evenness (E) Treatment Pre-treatment Post-treatment 22 25 15 0.686 2.34 1.79 0.76 0.56 Control Pre-treatment Post-treatment 22 25 15 0.751 2.10 1.97 0.68 0.61 Fourteen species of ant in five genera were detected at 20% of sites in any given year of the study (Table 11). These 14 species included representatives from all functional groups. We did not find that treatment impacted the occurrence of any species (all p > 0.5). Mean species richness of ants did not differ by treatment type (F2,13 = 1.03, p = 0.39), timing (F1,13 = 0.06, p = 0.82), or the interaction of timing and treatment (F2,13 = 1.66, p = 0.23). The only significant effect on species richness of ants was year (F2,13 = 3.85, p= 0.008), indicating that the ant species found on our study units experienced significant annual variation in site occupancy. Table 11. Ant species detected on study units in the Lake Tahoe Basin, 2006-2011. Species Ecological Group Campontus modoc Campontus vicinus Formica accreta Formica aserva Formica fusca Formica microphthalma Formica neoclara Formica neorufibaris Formica propinqua Formica ravida Formica sibylla Lasius pallitarsus Myrmica tahoensis Tapinoma sessile Decomposer Generalist Decomposer Decomposer Aerator Generalist Generalist Generalist Compiler Compiler Aerator Generalist Aerator Generalist 35 5.0 DISCUSSION 5.1 VEGETATION AND FUELS Our vegetation results indicated that treatment implementation, while varied, achieved the stated objectives. Implementation of fuels treatments resulted in more open forest with fewer smaller diameter trees. Pre-treatment forests were, with the exception of one east-shore treatment unit, dominated by firs (as measured by stems/acre), and had high densities of small (< 12” dbh) trees. Fuels treatments preferentially removed these smaller diameter trees, generally leaving forests dominated by moderatesized trees (12-24” dbh) trees. As a result of this preferential removal of smaller trees, average tree diameter increased on all plots following treatment. Post-treatment forests also had a greater proportion of pine than did pre-treatment forests, though forests remained fir-dominated. In addition to changes in live tree density, snag density was also reduced as a result of fuels treatments. This was not limited to small diameter snags, as moderate-sized snags were also less abundant on many units following treatment. Given the importance of snags as wildlife nesting and roosting sites, all treatment prescriptions set targets for snag retention. All treatment units met the prescription goals, which called for retention of at least 7.5 snags/ha. In addition to reductions in tree density and cover, forest understory cover was reduced on all treatment sites, significantly so on mechanically treated sites. Both measures of understory vegetation, shrub cover and herbaceous cover, declined following treatment. However, given the opening of forest canopies following treatment, additional light penetration would likely result in increases in both shrub and herbaceous cover over longer time periods. At this time, we have no data to allow us to predict the pace of this regeneration, or how it may vary based on treatment type or location. Our other measure of understory structure, coarse woody debris, was substantially reduced across all treatment units. The degree of change in forest structure varied greatly between treatment units. This was a result of several factors. By design, the treatment units had different diameter limits in treatment prescriptions. In addition, not all of our sampling units experienced treatment across the entire sampling area. Several sites had riparian buffer strips along stream channels, areas that were left untreated. These untreated areas create heterogeneity in post-treatment conditions within and across our survey units. Not surprisingly, the treatments altered forest structure in a manner consistent with the basic fuel reduction principles proposed by Agee and Skinner (2005): reduced tree density, especially in the smaller tree size classes, reduced ladder fuels (as indicated by canopy base height), and to a lesser extent reduced canopy fuels (Figures 16 and 17). However, the obvious piece missing with respect to the basic fuel reduction principles is reduced surface fuels. There was little evidence for a reduction of surface fuel loads across the treatments studied; in fact there was some evidence to the contrary, namely increased fuel depths in the masticated units (Table 5). Note that with the exception of increasing fuel depth in masticated units, the treatments did not noticeable increase fine woody fuel loads, indicating that the activity fuel treatments, namely piling and burning, were effective. Despite this general lack of change in surface fuels our fire behavior/effects modeling suggests that the modification of forest structure alone in the treatments studied did significantly alter tree survivability under wildfire conditions, particularly among the large trees. Based on the numerous studies demonstrating the importance of reducing surface fuels in reducing fire behavior and fire effects (Raymond and Peterson 2005, Stephens et al. 2009, Fulé et 36 al. 2012, Safford et al. 2012) it is reasonable to assume that if a surface fuel treatment component (e.g., prescribed burning) were added to the treatments studied tree survivability under wildfire conditions would only improve. Our findings indicating only a marginal effect of treatment intensity on predicted fire-induced tree mortality suggests that treatment effectiveness is not necessarily driven by the amount of basal area removed. This supports findings from a previous study demonstrating that differing upper diameter-limits for fuel reduction thinning had little effect on modeled landscape-level fire behavior (Collins et al. 2011). What is interesting about these results is that the reduction in density of small trees (<30.4 cm DBH), coupled with activity fuel treatment, alone can reduce large tree mortality due to wildfire. 5.2 WILDLIFE In spite of substantial changes to forest structure and composition following fuels treatments, we observed few significant changes in the diversity and abundance of wildlife. For small mammals, community composition experienced little change following treatment. Two new species were detected following treatment, while two species present before treatment were not detected after treatments. These species were all rare to uncommon during the time periods in which they were detected. Small mammal community evenness remained remarkably similar following treatment, as suggested by the minor changes to species composition. Our indices of change in the mammal community (Shannon’s Diversity and Evenness indices, Renkonen’s index) indicated that pre- and post-treatment communities were very similar in both species diversity and relative abundance of species. Small mammal abundance showed an overall positive trend following treatment implementation, driven primarily by an increase in chipmunk abundance. While none of these responses were statistically significant, mammal abundance was consistently higher in both treatment types and in both years following treatment implementation. Many of the components of forest structure that were reduced following treatment (e.g., shrub and herbaceous cover, coarse woody debris) are recognized as essential habitat features for many small mammals (Wilson and Carey 2000, Carey and Harrington 2001). It remains unclear which components of the fuels treatment are responsible for this positive trend, but the observed increase in overall abundance is consistent with the findings of several prior studies conducted elsewhere in the mountain West (Converse et al. 2006, Bagne and Finch 2010). At the species level, we observed significant responses to treatment in only two species. Deer mice populations declined by 67% on mechanical treatment plots in the first year following treatment, but returned to pre-treatment levels by the second year after treatment. The significant initial decline contrasts with the findings from several other studies which found increases in deer mouse abundance immediately following thinning in other parts of the western United States (Carey and Wilson 2001, Suzuki and Hayes 2003). This may be more reflective of response to the mastication which occurred on mechanically thinned plots, rather than a response to the mechanical thinning itself. Given the thick layer of masticated material and reduction in understory cover, this treatment type may have temporarily reduced habitat suitability for deer mice. Likewise, lodgepole chipmunks declined significantly on hand treatment plots in 37 the first year following treatment, but had returned to pre-treatment abundance by the second year following treatment. Although all other species-level results were non-significant, there were several consistent patterns observed. In particular, yellow-pine chipmunks (Tamias amoenus) increased in abundance on both mechanical and hand treatment units during both years of post-treatment surveys, while northern flying squirrels (Glaucomys sabrinus) declined on all treatment units during both years of the study. Finally, when considered as a group, chipmunks increased in abundance in both years following treatment on both hand and mechanical units. However, the only statistically significant increase was in the 2nd year posttreatment on mechanical sites. This lack of statistical significance for many results, in spite of large differences in abundance, suggests that wildlife abundance may be driven by other factors that overwhelm treatment effects. Like the small mammal community, the bird community at our study locations exhibited minor change following treatment. We detected 63 species total, 6 of which were detected only in pre-treatment samples, and 10 of which were found exclusively in post-treatment surveys. However, all of these species were rarely detected. As suggested by the small change in species richness, the bird community at our study site showed little change following treatments, as indicated by our indices of community diversity, similarity, and evenness. Overall abundance of birds showed a slight decline in both treatment types and in both years posttreatment. Bird abundance declined between 4.8 and 9.5 %, depending on year and treatment type. While this was a consistent pattern, none of these declines were statistically significant. When examined at the species level, we observed significant responses to treatment in 7 of the 36 species we analyzed. Goldencrowned Kinglet, Red-breasted Nuthatch, and Nashville Warbler all responded negatively to mechanical treatment in both years following treatment, with significant responses in at least one of the two posttreatment years. Kinglets and nuthatches are typically found in dense, shaded forest, so it is unsurprising that they were less common in the open forests found following treatment (Hayes et al. 2003). In addition, Nashville Warblers are found in forests with dense shrub cover, which they use for nest concealment. The reduction in shrub cover following treatment likely explains their decline. Four species exhibited significant positive responses to treatment. Red Crossbills responded positively to hand treatment in the first year post-treatment, but had returned to baseline levels by year 2. Similarly, Evening Grosbeaks showed a positive response to mechanical treatment only in the first year, suggesting that both of these irruptive finch species may have benefited from the disturbed conditions immediately following treatment. Dusky Flycatchers responded positively to hand thinning in the second year following treatment, possibly due to more open forest facilitating aerial foraging. Finally, Dark-eyed Juncos responded positively to mechanical treatments in both years post-treatment, perhaps a result of better ground foraging opportunities in the disturbed and exposed soil following treatment (Hayes et al. 2003). We observed significant results primarily in abundant species, indicating that we likely lacked power to detect significant responses to treatment in all but the most common species. Although we did not detect significant responses in most of the less common species, we did observe a number of consistent patterns 38 when considering response irrespective of statistical significance. However, these patterns were evenly split between species that responded positively and those that responded negatively to treatment. The treatments did not produce a pronounced effect in the bird community as a whole, as decreases in one species appear to have been mirrored by increases in another species. These decidedly mixed results are consistent with our community-level results, further indicating that while abundance of individual species may change significantly following treatment, the community as a whole experienced minimal change in species richness or evenness. Ants were not affected by either form of fuel reduction treatment utilized in this study. The majority of the ant species in our study nested in the ground, in logs and coarse woody debris, and in thatch mounds. Although prescriptions may reduce the amount of coarse woody debris in an area, an important habitat feature for foraging and nesting ants, we did not find a significant effect of hand or mechanical treatment on ant community composition or species richness. Additionally, all functional groups of ants were represented before and after treatment, indicating that ecosystem service provided by ants, including nutrient cycling and availability, soil aeration, water infiltration was maintained in treated forests (Folgarait 1998). The maintenance of species richness and composition suggests that upland forest treated by hand or mechanical means provide adequate habitat, at least initially, for this important invertebrate group. Differences in ant diversity may be short-term responses to disturbance of treatment. Treatment that disturbs soil and ground cover is likely to affect ground nesting and pile nesting ant species. Additionally, it appears that there was a year effect on ant diversity, as indicated by the decrease in diversity and evenness on control sites, although it was not as pronounced as declines on treated sites. It appears likely that year and treatment effects combine to affect community composition. Further study is needed to determine if changes in community composition are long-term or if diversity returns to pre-treatment levels over time. 6.0 CONCLUSION Our results indicate that conventional mechanical- and hand-thin fuels treatments are consistent with maintenance of diversity and evenness of ants, forest birds and small mammals. This is supported by other recent research suggesting minimal effects of fuels treatment at the community level (Stephens et al. 2012). Given the lack of longer-term research examining response to fuels treatments, we are unable to predict whether our short-term results would be consistent over longer, management-relevant time scales. At the species level, we did observe substantial changes in abundance for many species following treatment. However, in general our results were significant for only the most abundant species, which are typically not the drivers of land management decisions. This suggests that we lacked power to detect trends in all but the most abundant species, and that other variables, such as variation in treatment intensity or annual climatic variability, may have overwhelmed treatment response for many species. We hope to combine our results here with other ongoing SNPLMA-funded research to better assess specieslevel responses to fuels treatments. More broadly, we observed consistent, non-significant increases in overall small mammal abundance across both treatment types, and non-significant decreases in overall 39 bird abundance following treatment. The increase in small mammal abundance, while not statistically significant, is consistent with the results of several other studies (Converse et al. 2006, Stephens et al. 2012), and appears to be a widespread short-term response. The decrease in bird abundance has not been reported consistently by other authors, but we hypothesize that the slight decline observed here may be attributed to the decrease in structural diversity following treatment implementation. However, it is worth highlighting two caveats to our results. First, treatment intensity and extent varied substantially across treatment units, and the presence of untreated areas with treatment plots may have provided refugia for species dependent upon dense forested stands. Additionally, the short-term results observed here may not be predictive of longer-term trends in community diversity and evenness. For example, understory vegetation is likely to increase in extent following treatment, but did not do so over the time frame of our research. Changes in extent of shrub and herbaceous cover are likely to influence occupancy and abundance of several species of mammals and birds (Converse et al. 2006). However, we are unable to offer reliable predictions about response beyond the time frame of our research. Unfortunately, research examining longer-term wildlife response to fuels treatments is almost entirely lacking (Stephens et al. 2012), which leaves land managers without robust management information at relevant time scales. We hope that this prominent information gap will be addressed in future studies. 40 LITERATURE CITED Agee, J. K., and C. N. Skinner. 2005. Basic principles of forest fuel reduction treatments. Forest Ecology and Management 211:83–96. Agee, J. K. 2002. The fallacy of passive management: managing for firesafe forest reserves. Conservation in Practice 3:18–26. Andersen, A. 1997. Functional groups and patterns of organization in North American ant communities: a comparison with Australia. Journal of Biogeography 24:433–460. Bagne, K. E., and D. M. Finch. 2010. Response of small mammal populations to fuel treatment and precipitation in a ponderosa pine forest, New Mexico. Restoration Ecology 18:409–417. Barbour, M., E. Kelley, P. Maloney, D. Rizzo, E. Royce, and J. Fites-Kaufmann. 2002. Present and past old-growth forests of the Lake Tahoe Basin, Sierra Nevada, US. Journal of Vegetation Science 13:461–472. Beaty, R. M., and A. H. Taylor. 2009. A 14 000 year sedimentary charcoal record of fire from the northern Sierra Nevada, Lake Tahoe Basin, California, USA. The Holocene 19:347–358. Bestelmeyer, B. T., and J. A. Wiens. 1996. The effects of land use on the structure of ground-foraging ant communities in the Argentine Chaco. Ecological Applications 6:1225–1240. Bigelow, S. W., and P. N. Manley. 2009. Vegetation response to fuels management in the Lake Tahoe basin. Pages 41–82 in. Effects of fuels management in the Lake Tahoe basin: A scientific literature review. US Dept. of Agriculture, Forest Service, Pacific Southwest Research Station, Davis, CA. Brown, J. K., and C. M. Johnston. 1976. Debris Prediction System. Dept. of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. Brown, R. T. 2002. Thinning, fire and forest restoration: A science-based approach for national forests in the interior Northwest. Defenders of Wildlife, Washington, DC. Burnham, K. P., and D. R. Anderson. 2002. Model selection and multimodel inference: a practical information-theoretic approach. 2nd edition. Springer Verlag, New York. Carey, A. B., B. L. Biswell, and J. W. Witt. 1991. Methods for measuring populations of arboreal rodents. US Forest Service General Technical Report PNW-GTR-273. Carey, A. B., and C. A. Harrington. 2001. Small mammals in young forests: implications for management for sustainability. Forest Ecology and Management 154:289–309. Carey, A. B., and S. M. Wilson. 2001. Induced spatial heterogeneity in forest canopies: responses of small mammals. The Journal of Wildlife Management 1014–1027. Carey, H., and M. Schumann. 2003. Modifying wildfire behavior - The effectiveness of fuel treatments. National Community Forestry Center, Sante Fe, NM. Carlton, D. 2005. Fuel Management Analyst Plus v. 3.0. Fire Program Solutions, LLC., Estacada, OR, USA. Collins, B. M., S. L. Stephens, G. B. Roller, and J. J. Battles. 2011. Simulating fire and forest dynamics for a landscape fuel treatment project in the Sierra Nevada. Forest Science 57:77–88. Converse, S. J., G. C. White, K. L. Farris, and S. Zack. 2006. Small mammals and forest fuel reduction: national-scale responses to fire and fire surrogates. Ecological Applications 16:1717–1729. Dodson, E., and D. W. Peterson. 2009. Contributions of fire-killed trees to future wildlife habitat and surface fuels in dry coniferous forests. Logan, UT. Finney, M. A. 2001. Design of regular landscape fuel treatment patterns for modifying fire growth and behavior. Forest Science 47:219–228. Folgarait, P. J. 1998. Ant biodiversity and its relationship to ecosystem functioning: a review. Biodiversity and Conservation 7:1221–1244. Fulé, P. Z., J. E. Crouse, J. P. Roccaforte, and E. L. Kalies. 2012. Do thinning and/or burning treatments in western USA ponderosa or Jeffrey pine-dominated forests help restore natural fire behavior? Forest Ecology and Management 269:68–81. 41 George, T. L., and S. Zack. 2001. Spatial and temporal considerations in restoring habitat for wildlife. Restoration Ecology 9:272–279. Hayes, J. P., J. M. Weikel, and M. M. P. Huso. 2003. Response of birds to thinning young Douglas-fir forests. Ecological Applications 13:1222–1232. Knapp, E. E., J. M. Varner, M. D. Busse, C. N. Skinner, and C. J. Shestak. 2011. Behaviour and effects of prescribed fire in masticated fuelbeds. International Journal of Wildland Fire 20:932–945. Lutes, D. C. 2006. FIREMON: Fire effects monitoring and inventory system. US Dept. of Agriculture, Forest Service, Rocky Mountain Research Station. Manley, P., D. Murphy, L. Campbell, K. Heckmann, S. Merideth, S. Parks, M. Sanford, and M. Schlesinger. 2006. Biotic diversity interfaces with urbanization in the Lake Tahoe Basin. California Agriculture 60:59–64. Manley, P. N. 2009. Wildlife habitat and community responses to fuels management. Pages 224–302 in. Effects of fuels management in the Lake Tahoe basin: A scientific literature review. US Dept. of Agriculture, Forest Service, Pacific Southwest Research Station, Davis, CA. Murphy, D. D., and C. M. Knopp. 2000. Lake Tahoe Watershed Assessment. Volume 1. Pacific Southwest Research Station, USDA Forest Service, Albany, CA. Murphy, J., D. Johnson, W. Miller, R. Walker, E. Carroll, and R. Blank. 2006. Wildfire effects on soil nutrients and leaching in a Tahoe Basin watershed. Journal of Environmental Quality 35:479– 489. North, M., P. Stine, K. O’Hara, W. Zielinski, and S. Stephens. 2009. An ecosystem management strategy for Sierran mixed-conifer forests (General Technical Report PSW-GTR-220). Davis, CA: US Department of Agriculture, Forest Service. Omi, P. N., and K. D. Kalabokidis. 1991. Fire damage on extensively vs. intensively managed forest stands within the North Fork Fire, 1988. Northwest Science 65:149–157. Parsons, D. J., T. W. Swetnam, and N. L. Christensen. 1999. Uses and limitations of historical variability concepts in managing ecosystems. Ecological Applications 9:1177–1178. Peterson, D. L., M. C. Johnson, J. K. Agee, T. B. Jain, D. McKenzie, and E. D. Reinhardt. 2005. Forest structure and fire hazard in dry forests of the Western United States. GTR-PNW-628, US Dept. of Agriculture, Forest Service, Pacific Northwest Research Station, Portland, OR. Pielou, E. C. J. 1966. The measurement of diversity in different types of biological collections. Journal of theoretical biology 13:131–144. Pollet, J., and P. N. Omi. 2002. Effect of thinning and prescribed burning on crown fire severity in ponderosa pine forests. International Journal of Wildland Fire 11:1–10. Raymond, C. L., and D. L. Peterson. 2005. Fuel treatments alter the effects of wildfire in a mixedevergreen forest, Oregon, USA. Canadian Journal of Forest Research 35:2981–2995. Safford, H. D., J. T. Stevens, K. Merriam, M. D. Meyer, and A. M. Latimer. 2012. Fuel treatment effectiveness in California yellow pine and mixed conifer forests. Forest Ecology and Management 274:17–28. Sanford, M. P., P. N. Manley, and D. D. Murphy. 2009. Effects of urban development on ant communities: Implications for ecosystem services and management. Conservation Biology 23:131–141. SAS Institute. 2008. SAS 9.2. Cary, NC, USA. Schlesinger, M. D., P. N. Manley, and M. Holyoak. 2008. Distinguishing stressors acting on land bird communities in an urbanizing environment. Ecology 89:2302–2314. Shannon, C. E. 1949. Communication theory of secrecy systems. Bell System Technical Journal 28:656– 715. Stanton, A. E., and S. N. Dailey. 2007. Pre-treatment and partial-treatment forest structure and fuel loads in the Lake Tahoe Basin Management Unit. BMP Ecosciences, San Francisco, California. Stephens, S. L., J. D. McIver, R. E. J. Boerner, C. J. Fettig, J. B. Fontaine, B. R. Hartsough, P. L. Kennedy, and D. W. Schwilk. 2012. The Effects of Forest Fuel-Reduction Treatments in the United States. BioScience 62:549–560. 42 Stephens, S. L., J. J. Moghaddas, C. Edminster, C. E. Fiedler, S. Haase, M. Harrington, J. E. Keeley, E. E. Knapp, J. D. McIver, K. Metlen, C. N. Skinn, and A. Youngblood. 2009. Fire treatment effects on vegetation structure, fuels, and potential fire severity in western US forests. Ecological Applications 19:305–320. Stephens, S. L., and J. J. Moghaddas. 2005. Fuel treatment effects on snags and coarse woody debris in a Sierra Nevada mixed conifer forest. Forest Ecology and Management 214:53–64. Stephens, S. L., and L. W. Ruth. 2005. Federal forest-fire policy in the United States. Ecological Applications 15:532–542. Stephens, S. L. 1998. Evaluation of the effects of silvicultural and fuels treatments on potential fire behaviour in Sierra Nevada mixed-conifer forests. Forest Ecology and Management 105:21–35. Stephenson, N. L. 1999. Reference conditions for giant sequoia forest restoration: structure, process, and precision. Ecological Applications 9:1253–1265. Suzuki, N., and J. P. Hayes. 2003. Effects of thinning on small mammals in Oregon coastal forests. The Journal of Wildlife Management 352–371. Taylor, A. H. 2004. Identifying forest reference conditions on early cut-over lands, Lake Tahoe Basin, USA. Ecological Applications 14:1903–1920. Western Regional Climate Center. 2012. Tahoe, California - Climate Summary. <http://www.wrcc.dri.edu/cgi-bin/cliMAIN.pl?ca8758>. Accessed 24 May 2012. Wilson, S. M., and A. B. Carey. 2000. Legacy retention versus thinning: influences on small mammals. Northwest Science 74:131–145. Wolda, H. 1981. Similarity indices, sample size and diversity. Oecologia 50:296–302. 43 APPENDICES Appendix A. Vegetation Data Collection Detailed Protocol Vegetation was measured within a fixed 17.58 meter (58 ft) radius plot of 0.1 ha (0.25 ac). A random azimuth was selected for the first transect and then three additional transects were established at subsequent 90° angles with the 0 meter marks at the plot center. The four transects were the basis for sampling down woody debris (DWD), herbaceous and shrub cover and frequency, and duff and litter depths (Figure A-1). Plots were monumented with five 2 foot pieces of rebar capped with yellow plastic caps imprinted with “USFS UPFU”; one at the plot center and one at the distal end of each transect. Vegetation Plot Layout Transect 1 Transect 4 Transect 2 Transect 3 Tree Macroplot 17.58 m (57.68’) radius Sapling and Seedling Subplot 3.57 m (11.77’) radius- tree saplings and seedlings less than 1 cm dbh. DWD 17.58 m transects, 0 is at plot center. 1 and 10hr fuels counted from 15-17m, 100 hr from 12-17 m, and 1,000 hr for the entire length. Litter and duff measurement and vegetation cylinder conducted at 8 m and 16 m. Cover and Frequency - .5 m2 quadrats on all transects at the 3, 6, 9, 12, and 15 m marks placed on the right side of transect (from center of plot). Permanent rebar stake locations- photo points are from outer 4 rebar stakes looking in to plot. Figure A1. Uplands Fuels research project vegetation plot lay-out. Plot Description Descriptive data collected in each plot included: UTM coordinate in NAD 27, slope, aspect, general landform, horizontal and vertical slope shape. The two most dominant species with greater than 10% canopy cover were recorded for three stratums; upper (> 3 m tall), mid (3 to 10 m tall), and low (< 1 m tall). A photo point was established at the distal end of each transect looking down the meter tape toward the plot center. Tree Data A breakpoint of 15 cm (6 in) diameter at breast height (DBH) was selected for sampling mature trees. The following data was recorded for all mature trees within the entire plot: species, DBH, total height, height to live crown base, live crown ratio, crown position, and observed damage. Snags were also sampled within the entire plot. Snag data included: species, DBH, total height, and decay class. Within a 3.57 m 44 (11.7 ft) fixed radius subplot of 0.01 ac (0.004ha) nested at the plot center, the species, DBH, total height, and live crown ratio was recorded for all saplings greater than 1.37 m (4.5 ft) tall with a DBH less than the breakpoint. Saplings were categorized by 4 size classes based on DBH: 0-2.5, 2.5-5, 5-10, 10-15 cm. The number of all seedlings less than of 1.37 meters tall was recorded for each species. Seedlings were categorized by 5 height classes: 1-15, 15-30, 30-60, 60-100, and 100-140 cm. The midpoint values of size and height classes were used in calculations. Each mature tree and snag was permanently marked with an aluminum tree tag and nail. Canopy cover was measured at 25 points using a 5 m by 5 m grid using a GRS site-tube densitometer. Fuels Data Surface and ground fuels were sampled on all four transects in each inventory plot using the line-intercept method (Brown, 1974). One-hour (0-0.64 cm) and ten-hour (0.64-2.54 cm) fuels were tallied from 15-17 meters, 100-hour (2.54-7.62 cm) fuels from 12-17 meters, and 1000-hour (>7.62 cm) fuels were sampled along the entire length (17.84 meters) of each transect. The larger fuels (1000-hour) represent coarse woody debris (CWD) that has high value for many wildlife species, so the following information was collected for each CWD: species, diameter at the tape, diameter at each end, length, and decay class. Duff and litter depth (cm) was measured at the 8 and 16 meter marks. At the same locations, the surveyors estimated the following within an imaginary 2m by 2m cylinder: live and dead tree/shrub cover, average tree/shrub height, live and dead herb cover, and average herb height. Measurements were conducted towards the distal ends of the transects to avoid the disturbance that was generally concentrated in the plot center. Herbaceous and Shrub Cover/Frequency Herb and shrub percent cover, height, and nested frequency were measured in five 0.25m2 quadrats located at 3 meter intervals (3,6,9,12,15) along all four transects, for a total sample area of 1.25 m2. Frequency describes the abundance and distribution of species and is very useful for comparing significant differences between two plant communities or detecting significant change in a single community over time. A reasonable sensitivity to change results from capturing frequencies between 20 and 80 percent, and therefore, a nested quadrat system was used to avoid problems resulting from using a single quadrat size. Nested quadrat sizes (5x5, 25x25, 25x50, 50x50 cm) corresponded to a nested rooted frequency ratio of 1:25:50:100. Plant cover was measured as the vertical projection of foliage within a percentage of the quadrat and the percent value indicates the relative influence of each species on the community. A system of 12 cover classes, (0-1, 1-5, 5-15, 15-25, 25-35, 35-45, 45-55, 55-65, 65-75, 75-85, 85-95, 95-100) was used to reduce human error and increase the consistency of estimates. Midpoint values were used for computation. Height gives detailed information about the vertical distribution of plant species cover with in the plot. It allows calculations of 1) plant species volume (cover x height) and 2) biomass (height x cover x bulk density). FIREMON uses 0.8 kg/ m3 for herbaceous BD and 1.8 kg/m3 for shrubs. 45 Appendix B. Total number of individuals and site frequency of small mammal species caught and marked during live-trapping surveys conducted late May through August across 16 units in the Lake Tahoe Basin, 2006-2011. Species are ranked in order of abundance. Common Name Deer mouse Long-eared chipmunk Allen’s chipmunk Yellow-pine chipmunk Lodgepole chipmunk Golden-mantled ground squirrel Douglas’ squirrel Northern flying squirrel California ground squirrel Trowbridge’s shrew Least chipmunk Long-tailed vole Pinyon mouse Vagrant shrew Brush mouse Bushy-tailed woodrat Scientific Name Peromyscus maniculatus Tamias quadrimaculatus Tamias senex Tamias amoenus Tamias speciosus Spermophilus lateralis Tamiasciurus douglasii Glaucomys sabrinus Spermophilus beecheyi Sorex trowbridgii Tamias minimus Microtus longicaudus Peromyscus truei Sorex vagrans Peromyscus boylii Neotoma cinerea 46 Total Individuals 811 611 507 494 487 312 137 118 64 23 19 16 10 9 3 2 Site Count 16 16 16 12 10 8 14 16 9 8 7 7 6 4 2 2 Appendix C. Total number of detections and site frequency of all bird species detected during point count surveys conducted late May through early July across 16 units in the Lake Tahoe Basin, 2006-2011, ranked in order of abundance. Common Name Mountain Chickadee Red-breasted Nuthatch Yellow-rumped Warbler Golden-crowned Kinglet Dark-eyed Junco Western Tanager Dusky Flycatcher Evening Grosbeak Steller's Jay Fox Sparrow Nashville Warbler Brown Creeper American Robin Hermit Thrush Warbling Vireo Pine Siskin Brown-headed Cowbird Western Wood-Pewee White-headed Woodpecker Cassin's Finch Townsend's Solitaire Hairy Woodpecker Hermit Warbler MacGillivray's Warbler White-breasted Nuthatch Red Crossbill Cassin's Vireo Clark's Nutcracker Northern Flicker Olive-sided Flycatcher Red-breasted Sapsucker Williamson's Sapsucker Chipping Sparrow Black-backed Woodpecker Band-tailed Pigeon Ruby-crowned Kinglet Pygmy Nuthatch Wilson's Warbler Scientific Name Poecile gambeli Sitta canadensis Dendroica coronata Regulus satrapa Junco hyemalis Piranga ludoviciana Empidonax oberholseri Coccothraustes vespertinus Cyanocitta stelleri Passerella iliaca Oreothlypis ruficapilla Certhia americana Turdus migratorius Catharus guttatus Vireo gilvus Carduelis pinus Molothrus ater Contopus sordidulus Picoides albolarvatus Carpodacus cassinii Myadestes townsendi Picoides villosus Dendroica occidentalis Oporornis tolmiei Sitta carolinensis Loxia curvirostra Vireo cassinii Nucifraga columbiana Colaptes auratus Contopus cooperi Sphyrapicus ruber Sphyrapicus thyroideus Spizella passerina Picoides arcticus Columba fasciata Regulus calendula Sitta pygmaea Wilsonia pusilla 47 Total Detections 1244 964 848 732 689 613 575 551 537 534 264 263 259 223 201 172 161 148 145 129 121 105 105 89 79 70 49 41 40 29 28 26 25 24 24 21 19 18 Site Count 16 16 16 16 16 16 16 16 16 16 15 16 16 16 13 16 12 16 16 15 15 15 12 15 13 9 7 4 12 9 13 7 9 9 7 2 6 5 Common Name Pileated Woodpecker Common Raven Green-tailed Towhee Mountain Quail Brewer's Blackbird Black-headed Grosbeak House Wren Mourning Dove Hammond's Flycatcher Calliope Hummingbird Sooty Grouse Orange-crowned Warbler Red-shouldered Hawk Cooper's Hawk Black-throated Gray Warbler European Starling Golden-crowned Sparrow Red-tailed Hawk Red-winged Blackbird Spotted Sandpiper Rufous Hummingbird Sharp-shinned Hawk Northern Goshawk Pine Grosbeak Pacific Wren Bewick's Wren Downy Woodpecker Great Horned Owl Lincoln's Sparrow Osprey Song Sparrow Townsend's Warbler Western Bluebird Yellow Warbler Scientific Name Dryocopus pileatus Corvus corax Pipilo chlorurus Oreortyx pictus Euphagus cyanocephalus Pheucticus melanocephalus Troglodytes aedon Zenaida macroura Empidonax hammondii Stellula calliope Dendragapus fuliginosus Vermivora celata Buteo lineatus Accipiter cooperii Dendroica nigrescens Sturnus vulgaris Zonotrichia atricapilla Buteo jamaicensis Agelaius phoeniceus Actitis macularia Selasphorus rufus Accipiter striatus Accipiter gentilis Pinicola enucleator Troglodytes pacificus Thryomanes bewickii Picoides pubescens Bubo virginianus Melospiza lincolnii Pandion haliaetus Melospiza melodia Dendroica townsendi Sialia mexicana Dendroica petechia 48 Total Detections 17 15 9 8 7 7 7 7 6 6 6 4 4 4 3 3 3 3 3 3 2 2 2 2 2 1 1 1 1 1 1 1 1 1 Site Count 4 5 4 7 6 5 3 3 4 3 3 3 3 2 2 2 2 2 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1