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ASSESSING RESTORATION SUCCESS FOR A WET MONTANE

SIERRA NEVADA MEADOW

______

A Thesis

Presented

to the Faculty of

California State University, Chico

______

In Partial Fulfillment

of the Requirements for the Degree

Master of Science

Biological Sciences

______

Rachel Schleiger

Spring 2018

TABLE OF CONTENTS

PAGE

List of Tables ...... v

List of Figures ...... vii

Abstract ...... viii

CHAPTER

I. Introduction ...... 1

Extent of Damage and Motivation for Restoration ...... 1 Restoration Definition and Goals...... 2 Trajectory Models ...... 4 Study Objectives and Hypothesis ...... 5

II. Methodology ...... 7

Study System ...... 7 Field Sites...... 9 Vegetation Sampling ...... 12 Vegetation Mapping...... 14 Vegetation Characterization...... 15 Soil Moisture ...... 16 Data Analysis and Statistics ...... 18

III. Results ...... 22

Variation Between Watersheds and Between Sampling Years ...... 22 Vegetation and Soil Moisture ...... 23 Richness ...... 37 Invasive and Non-Native Species ...... 47 Summation of Important Points ...... 54

IV. Discussion ...... 57

Disturbance in Wet Montane Meadows ...... 57 Comparison Issues with Less-Disturbed Sites ...... 58 Summary of Key Findings ...... 58 Restoration Assessment ...... 59

iii References ...... 64

Appendices

A. Plant List ...... 74 B. USFWS Indicator Categories ...... 80 C. Patch Classification and Characteristics Across Sites for 2012 Field Season ...... 82 D. Number of Patches at Each Site Dominated by Different Plant Types ...... 89 E. Areas (m2) of Plant Type (a) and Plant Wetland Affinities (b) by Soil Moisture Categories for each Site in 2012 Field Season ...... 91 F. Percentage of Site Area in Each Soil Moisture Category and Total Area of Non-vegetated (NV) Ground (m2) and the (Percent Area that Comprises each Moisture Category) for 2012...... 94

iv LIST OF TABLES

TABLE PAGE

1. Site Locations within Watersheds (Separated by Different Hydrologic Unit Scales) with Unit Name, Resolution, and Area ...... 11

2. Total Precipitation Averages (inches), using the Current Water Year System, between Watersheds for Both Sampling Years ...... 22

3. Total Estimated Meadow Area (m2) and Number of Patches and (Vegetation Patch Types) Surveyed for 2012 ...... 24

4. Study Sites Classified by 2012 Patch Soil Moisture Categories...... 25

5. Richness and (Percent Total Richness) for Native, Invasive, and Non-native Species as well as Total Species Richness and Family Richness Across Both Sampling Seasons ...... 38

6. Species Richness for Each Site Between Sampling Years (2011 and 2012) ...... 39

7. Percent of Meadow Area in Each Soil Moisture Category and the Accompanying Species Richness in 2012 for Each Site ...... 40

8. Species Richness and (Average Percent of Total Area) of Plant Types Across Sampling Years ...... 41

9. The Average Percentage of Total Area in Each Moisture Category Across Occupied by a Particular Plant Type in 2012 for Each Site ...... 43

10. Species Richness and (Average Percent of Total Area) Categorized According to the USGS Wetland Classification Categories Across Sampling Years ...... 44

11. Average Percent of Total Area within Each Site’s 2012 Patch Moisture Categories Occupied by Plant Species Classified by the USGS Wetland Classification System ...... 45

v TABLE PAGE

12. The Average Area of Non-vegetated Ground (m2) and (Average Percentage of Total Area) at Site Scale Between Years ...... 47

13. Average Area (m2) and (Average Percent of Total Area) Occupied by Invasive, Non-native, and Native Species for Each Sampling Year at Each Site ...... 49

14. The Number of Invasive and Non-native species by Plant Type and (Percent of Total Species Richness for Each Plant Type) at Each Site ...... 50

15. Invasive and Non-native Species Counts and (Percent of Total Species Richness at Each Site in the USDA Wetland Classification Categories) Across Both Sampling Seasons ...... 51

16. Average Area (m2) and (Average Percent of Total Area (%)) Across Both Sampling Seasons Occupied by the Most Prevalent Invasive and Non-native Species for Each Meadow Organized by their Wetness Affinities ...... 53

17. Table 17a-b. Relative Area for Each of the Moisture Categories in Each Site, Total Average Area (m2) and The (Average Percent of Total Area for Each Moisture Patch Occupied) by Invasive (a) and Non-native (b) Species Across the 2012 Sampling Season ...... 55

vi LIST OF FIGURES

FIGURE PAGE

1. Example Trajectory Model Illustrating Movement of Degraded State Through Time to Multiple Possible Endpoints Based on Level of Complexity and Function ...... 4

2. Restored meadow, Calaveras Big Trees State Park ...... 10

3. Position of all Field Sites in the San Joaquin River Regional Watershed, as well as its Smaller Bydrologic Units ...... 12

4. Position of Dorrington (DR), Hazel Fisher (HF), and Big Trees (BT) Study Sites in the Upper Calaveras and Upper San Antonio Creek Watersheds ...... 13

5. Position of El Capitan (EL) and Half Dome (HD) Study Sites in the Upper Merced as well as Tenaya Creek and Indian Canyon Creek- Merced River Watersheds ...... 14

6. Total Relative Area of Patches Classified by 2012 Soil Moisture Categories for Each Site ...... 26

7. Figures 7a-e: Average Soil Moisture Maps Across the Sampling Season, with Inset for June (Wettest Sampling Month) Moisture, for the Delineated Vegetation Patches Based on Field Sampling in June 2012 at Each Site: (a.) DR, (b.) HF, (c.) BT, (d.) EL, (e.) HD ...... 27

8. Figure 8a-e. Dominant Plant Type, or not, Across Both Sample Seasons at (a.) DR, (b.) HF, (c.) BT, (d.) EL, and (e.) HD ...... 32

9. Average Percent Non-vegetated Total Area Across Moisture Categories at Each Site for 2012 ...... 49

vii

ABSTRACT

ASSESSING RESTORATION SUCCESS FOR A WET MONTANE

SIERRA NEVADA MEADOW

Rachel Schleiger

Master of Science in Biological Sciences

California State University, Chico

Spring 2018

Meadows in the Sierra Nevada are characterized as wet, heterogeneous habitats with diverse plant communities, often being biodiversity hot spots. These meadows not only provide resources for wildlife but also filter and store snowmelt, providing sustained water sources for both wildlife and Californians. Recognition of meadow significance combined with persistent human disturbance motivates restoration efforts to improve hydrologic connections and biotic health within these meadows. This research is evaluating the trajectory of a restored montane meadow in Sierra Nevada, California. Comparing soil moisture, plant community composition, and exotic species extent of this restored meadow to disturbed and less disturbed meadows provided context for this assessment.

Results indicated soil moisture was highest in the restored meadow followed by the disturbed then less-disturbed sites. After further investigation, the less-disturbed sites were found to have significantly less total annual precipitation which greatly impacted overall plant composition. As such, the less-disturbed sites were deemed inappropriate for

viii

comparison and the restored sites trajectory assessment was thus more focused on comparisons to the disturbed sites. The restored site was found to have lower moisture heterogeneity but higher hydrologic connectivity compared to the disturbed sites, traits more definitive of wet montane meadows. Species richness, status, plant type, and wetland classifications at the restored site were also more definitive of wet montane meadows compared to the disturbed sites. However, the restored site did have concerning areas of exotic species, especially in drier soils. Currently, rewetting techniques applied appear to be successful. Although, it is clear that adaptive management is needed to address issues of concern and help keep a continued positive ecological trajectory at this restored site, especially when heading into an uncertain future.

ix 1

CHAPTER I

INTRODUCTION

Extent of Damage and Motivation for Restoration

The great extent of anthropogenic influence on atmospheric, marine, and terrestrial systems is causing pervasive damage to the structure, function, and biodiversity of all Earth’s ecosystems (Vitousek et al. 1997). These changes will likely be exacerbated by climate change (Society of Ecological Restoration (SER) 2004, Cortina et al. 2006). Repair or restoration of degraded ecosystems is seen as an important means of reversing this trend

(Hobbs and Harris 2001, Chapman and Underwood 2010).

Wetland systems, which support unique ecosystem services and biodiversity, are one of the most impacted habitat types across the globe (Purdy and Moyle 2006). For example, wetland meadows present unique and diverse environmental conditions compared to the surrounding landscape (Ratliff 1982). Globally, wet meadows have a history of human use largely centered on agriculture (Quetier et al. 2007, Zhao and Zhau 1999, Saberwal 1996,

Marini et al. 2007, Rochefort and Gibbons 1992, Austin et al. 2007, Hammersmark et al.

2008). Although agriculture is the root of most degradation issues, both historically and in modernity, intense recreational activities have started to become a concern for wetland meadows (Rochefort and Gibbons 1992, Egan et al. 2000, 2004). Variables such as intensity of human impacts and population density and length of human use, impact the level of degradation to wetlands (Sanderson et al. 2002). In many instances, there will always be human use and any attempts at restoration are compromised by human necessity (Saberwal

1996, Quetier et al. 2007). In some instances, however, human use has ceased and there have been attempts at restoration (Rochefort and Gibbons 1992, Hammersmark et al. 2008).

Assessments of wet meadow degradation have found that intense human usage has led to changes in both abiotic factors such as hydrology and soil nutrient composition

(Castelli et al. 2000, Quetier et al. 2007, Hammersmark et al. 2008), as well as biotic factors such as species richness (Marini et al. 2007, Freitas et al. 2014), composition (Austin et al.

2007, Lang and Halpern 2007), and biomass (Pope et al. 2015). Hydrologic disturbance has also led to meadow establishment by exotic species (Underwood et al. 2004, Loheide and

Gorelick 2005) and native shrub (Berlow et al. 2002) and conifer encroachment (Vale 1987,

Lang and Halpern 2007). With increased moisture gradients due to hydrologic damage, and higher incidence of exotic and woody species, disturbed sites could be observed with more species than their less-disturbed counterparts (Loheide and Gorelick 2005). As time propels these systems into an unpredictable future, especially regarding climate, there could be even more critical changes in the abiotic and biotic patterns of these meadows (Cortina et al. 2006,

Choi et al. 2008). Recognition of wetland meadows’ importance, often combined with a history of human disturbance, provides the opportunity for restoration efforts to improve the outlook for these wetland habitats. This study will focus on assessing a restored wetland meadow in Big Trees State Park, California, USA.

Restoration Definition and Goals

The science of restoration ecology and the practice of ecological restoration combined provide the required foundations for humanity to mitigate, prevent, and reverse ecosystem deterioration (SER 2004). As ecological restoration is applied based on subjective restoration goals, restoration definitions can vary (Jackson et al. 1995). Despite this variation,

3 all definitions incorporate some idea of stopping and reversing ecosystem damage with major human intervention. A 2006 definition presented by SER and the International Union for the

Conservation of Nature (IUCN), states, “Ecological restoration is the process of assisting the recovery of an ecosystem that has been degraded, damaged or destroyed.” Although broad, this definition implies that restoration requires serious practical obligations to actively engage and intervene in current social and environmental affairs (Andel and Aronson 2005).

This definition does not specify, as many original definitions did, on ecosystem recovery focused on some historical state. Most current papers agree that focus on historical states in restoration goals is impossible to achieve and inappropriate given that systems are ever in flux (Hobbs and Harris 2001, Falk et al. 2006, Choi 2007, 2008), especially considering predicted climate changes (Choi 2004, 2008, Harris et al. 2006).

Typical scientific-based goals of restoration are focused on recovering a natural range of ecosystem composition, structure, function, and dynamics to create a self-supporting ecosystem that is resilient to natural disturbance regimes (Falk 1990, SER 2004). However, restoration goals all vary based on contextual constraints and societal objectives, such as co- opting ecosystem services or modifying ecological attributes for human use (Falk et al.

2006). Thus, goals set for restoration initiatives do not always have a scientific base.

Nonetheless, many agree that goals need to be set that are not only ecologically achievable but also economically feasible and socially acceptable (Choi, 2004, 2007, Halle, 2007,

Hobbs, 2007).

Trajectory Models

Ecological theories have provided much of the conceptual frameworks for projecting future restoration outcomes. In particular, community assembly based on

4 successional models has been applied to restoration trajectories since the early twentieth century (Clements 1916, 1936, Gleason 1917, 1926). However, the current view of these trajectories is based on a newer successional model that asserts a degraded system can travel along several different (and unpredictable) trajectories towards some stable state (Figure 1)

(Sutherland 1974, Hobbs and Mooney 1993, Temperton et al. 2004). Unpredictable courses of community succession in an unforeseeable future environment add even more uncertainty to restoration trajectories (Matthews et al. 2009, Choi 2008). One also needs to consider the past history of the system in question, as it can have a large impact on the potential for restoration to reach the desired state (Hobbs and Mooney 1993). In this study, ecological trajectory concepts were utilized to create a heuristic hypothesis for restoration assessment.

Figure 1. Example trajectory model illustrating movement of degraded state through time to multiple possible endpoints based on level of complexity and function. Adapted from Hobbs and Mooney 1993.

Study Objectives and Hypothesis

The objective of the study was to assess the ecological trajectory of a restored wet meadow in Big Trees State Park, California, USA. The main goal for the restored wetland meadow assessed in this study was to refill the historic channeling that occurred, to raise the water table, and provide conditions supportive of native wet meadow plants. Thus, this restoration was focused on the Field of Dreams assumption that systems will self-organize based on restored abiotic conditions (Hilderbrand et al. 2005). Although this assumption generally does not follow basic ecological principals, in this habitat, where abiotic patterns drive biotic patterns, a positive outcome could still result from purely abiotic restoration.

Restoration goals that aim toward the future should also acknowledge the changing and unpredictable climate and how it will affect the dynamic nature of ecological communities with multiple potential trajectories (Choi 2008).

It was hypothesized that the ecological health of the wet meadow will have moved away from the “degraded state” (Hobbs and Mooney 1993) based on restoration techniques applied. Ecological health was assessed by making comparisons of ecological attributes of wetland meadows across a disturbance gradient. Attributes measured were species richness and percent relative foliar cover of native, non-native, and invasive plants and bare ground. Community composition was assessed relative to patch type (delineated dominant vegetation types), meadow condition, year, and wetness. Ultimately the apparent ecological trajectory will indicate if the restoration is complete, and thus whether or not the methods of restoration were successful.

CHAPTER II

METHODOLOGY

Study System

The Sierra Nevada from north to south is 640 km long, 105 km wide, and encompasses 63,100 km2 both in California and Nevada, USA. The foothills of the Sierra

Nevada start around 300m and rise to over 4,300 m in its peaks (McNab and Avers 1996).

There are many local climates due to diverse topography, elevation, and geographic position in relation to the Great Valley, Coast Ranges, and Pacific Ocean (Storer et al. 2004).

Temperature averages in summer range from 8.6 to 20.3 °C and 1.7 to -10.8 °C through the winter (McNab and Avers 1996, Storer et al. 2004). The bedrock is dominated by granite, which supports wet habitats with a neutral to slightly acidic pH runoff during the growing season, which ranges from 20-230 days depending on local conditions (McNab and Avers

1996, Storer et al. 2004). More than 50% of the total precipitation in the Sierra Nevada falls in January, February, and March, while less than 3% falls during summer months (McNab and Avers 1996). The snow line begins at elevations around 600-900m though elevations above 1800m where most snow falls (Storer et al. 2004). Generalized plant community belts follow elevational contours (foothill, mixed conifer, boreal, Jeffery pine, sagebrush, southeast desert) (McNab and Avers 1996, Storer et al. 2004).

Meadows represent roughly 10% of total area in the Sierra Nevada and are defined by their hydrology, vegetation, and soil characteristics (Ratliff 1982, 1985, Loheide et al. 2009). Meadows occur in areas with roughly flat landforms surrounded by steep

7 topographic contours, and large watersheds that offer a shallow water table and fine textured soils (Ratliff 1982, 1985). Meadow elevation also influences the types of vegetation that occur due to differences in growing season length, climate, and soil development. For example, montane meadows are surrounded by forest (915-2400m), subalpine meadows are near the upper limit of trees (2400-2900m), and alpine meadows are above the tree line and are generally surrounded by rocks and boulders (>2900m) (Ratliff 1985, Purdy and Moyle

2006). Sierra Nevada wet-meadow vegetation relies on shallow groundwater or surface water supplied by snowmelt during the growing season. Water tables in these meadows vary both spatially and temporally (Castelli et al. 2000). For these reasons, wet meadows are classified as hydrologic-dependent ecosystems (Ratliff 1985, Murray et al. 2003, Boulton 2005). Most meadows contain complex patterns of wet, moist, and dry areas that support one to two distinct plant community types based on the moisture mosaic (Sawyer, Wolf, and Evans

2009). These plant communities are dominated by herbaceous species while woody vegetation, like trees or shrubs, occur in low abundance due to high water tables.

Within individual meadows and across meadow types, plant species diversity is relatively high (Ratliff 1985, Debinski et al. 2000), and wet meadows represent diversity hot spots in the Sierra Nevada (Viers et al. 2013). Generally, sedges and rushes dominate meadow vegetation and those with stream banks are often lined with willows and alders

(Sawyer, Wolf, and Evans 2009). Sedges have long and dense root and rhizome networks that produce a sod that is inherently resistant to erosion (Purdy and Moyle 2006, Moyle et al.

2008). These networks of root mats act like sponges storing and keeping water available through much of the growing season (Ratliff 1985, Purdy and Moyle 2006, Moyle et al.

2008). Wetland meadows also provide important breeding grounds for invertebrates, which

8 are a key food source for many birds, amphibians, and reptiles (Ratliff 1985). Meadow vegetation also provides food and habitat structure for small mammals which, in turn, provide an important prey base for raptors, coyotes, and other predators (Ratliff 1982, 1985).

Field Sites

The focus wet meadow in this study, called the North Grove meadow (BT), is located within Big Trees State Park in Calaveras County, CA, USA (38°16’30.64”N

120°18’29.17”W) (Figure 2). At an elevation approximately 1422m (montane), BT is roughly oval shaped, positioned northeast to southwest with a 2% slope, and is slightly over

20,000m2. Before human influence (<1850s), BT was fed by an ephemeral stream that fanned out across most of the meadow creating hydric and mesic habitats during the growing season

(Boyd and Woodward 1988). However, as soon as Big Trees State Park’s location became popular (1852), the wet meadow was channelized to support a dairy, farm, and hotel. It was not until just after the parks establishment in 1931 that the agricultural practices were abandoned and the hotel was taken down due to heavy damage sustained through many winters (California State Parks 2004). After this, damage to the meadow was limited to human recreational use, such as trails and camping along its borders.

A 1988 vegetative assessment found that the hydrologic alterations

(channelization) not only severely lowered the water table by creating more xeric habitat conditions, but also dramatically depleted populations of native wet-meadow species dependent on hydric and mesic habitats (Boyd and Woodward 1988). Hence, the overall species richness and composition did not reflect a healthy meadow condition. To both increase the likelihood of native wet meadow species expansion and decrease the expansion of non-native, invasive, and woody species it was suggested to restore the hydrology (Boyd

Figure 2. Restored meadow, Calaveras Big Trees State Park. Retrieved from Google Earth. 2012, June. Calaveras Big Trees State Park. 38°”53”N 120°18”33”W. https://earth.google.com/web/

and Woodward 1988). In 2003, state park biologists implemented a restoration project to fill in the channelization and thus raise the water table to provide conditions supportive of marsh-meadow plants. By 2004 the project was complete, and the meadow had approximately seven years to regenerate populations of marsh-meadow plants before the start of this study in 2011.

To assess the ecological trajectory for this Big Trees restored meadow, an effort was made to select reference montane meadow sites along a hydrological disturbance gradient that had comparable ecological attributes. These attributes included characteristics that both define montane meadows as well as other features more unique to the restored meadow. Accordingly, reference meadows were found with the following attributes: seasonally wet between the months of May-September, an elevation around 1400m, overlain

10 on granite bedrock, aspect N-NW, within 0.25mi of a trail or road, surrounded by coniferous forest and near small groves of giant sequoias, and situated in the uppermost portion (largest hydrologic unit) of the watershed (Table1).

Table 1. Site locations within watersheds (separated by different hydrologic unit scales) with unit name, resolution, and area.

Resolutiona HUC Nameb Sites Within Area (km2) Upper San Antonio D, HF, BT 72.5 Creek 12digit-HU Tenaya Creek HD 80.3 Indian Canyon Creek- EL 80.3 Merced River Upper Calaveras D, HF, BT 992 8digit-HU Upper Merced EL, HD 2797.2 4digit-HU San Joaquin D, HF, BT, EL, HD 40403.8 aSize of the hydrologic unit (HU) code which reflects the area of the map layer polygon bHydrologic unit code (HUC) name associated with the map layer polygon

Using the above attributes, comparable meadows were identified then classified as either disturbed (channelization present) or less-disturbed (no channelization present).

Overall, two disturbed and two less-disturbed meadows were located within the San Joaquin regional watershed (Figure 3). The two hydrologically disturbed meadows were located within Stanislaus National Forest; with Dorrington roughly 7 km east of the restored meadow

(designated DR, 38°18’18.83”N 120°16’46.44”W) and Hazel Fisher roughly 7 km west

(designated HF, 38°15’39.18”N 120°20’37.82”W) (Figure 4). As such, both the restored and disturbed sites are situated in the same 12-digit hydrologic unit (Table 1).

There were no meadows located in the Upper Calaveras watershed that had the required ecological attributes and could also be classified with less hydrological disturbance.

As a result, the search for less-disturbed sites, still having all required ecological attributes, was expanded. Less-disturbed meadows were located in Yosemite National Park. The Half

Figure 3. Position of all field sites in the San Joaquin River regional watershed, as well as its smaller bydrologic units.

Dome site (designated HD, 37°45’14.40”N 119°32’39.69”W) is situated in Tenaya Creek while the El Capitan site (designated El, 37°43’21.47”N 119°38’48.09”W) is in Indian

Canyon Creek-Merced River portion of the Upper Merced watershed in Mariposa County

(Figure 5).

Vegetation Sampling

To compare vegetation of restored meadow (BT) to disturbed and less-disturbed sites, vegetation was sampled in 2011 and 2012 using a stratified random sample design.

Meadows were stratified into patches based on vegetation types using the CNPS definition as visually distinct and repeated entities (Sawyer, Wolf, and Evens 2009). To characterize

Figure 4. Position of Dorrington (DR), Hazel Fisher (HF), and Big Trees (BT) study sites in the Upper Calaveras and Upper San Antonio Creek watersheds. Study site outlines not to scale.

diversity within each patch, 0.5m2 sample plots were randomly assigned with respect to direction and haphazardly tossed relative to distance and angle. This random and haphazard strategy was used to account for observed increased variability of vegetation composition with patch size. The number of plots placed in each patch was determined by both size and heterogeneity of a patch. Thus, actual counts of plots placed in each patch from month to month and season to season were based on visual estimations of species accumulation curves, whereas if after 5 consecutive plots no new species were found, sampling ceased in that patch. Within each plot, the percent cover of each species present was visually estimated.

Typically transects are used in settings like these meadows, as was completed in the pre- restoration assessment of the focus meadow. However, with how heterogeneous this habitat is, high intensity transect sampling would be needed to get a clear picture of the vegetation

Figure 5. Position of El Capitan (EL) and Half Dome (HD) study sites in the Upper Merced as well as Tenaya Creek and Indian Canyon Creek-Merced River watersheds. Study site outlines not to scale.

patterning. The methods utilized in this study allowed for a more efficient means of sampling, meaning fewer samples needed to represent each patch, so as to least disturb the habitat during growing seasons sampled (2011 and 2012). All meadows were sampled during a one week period during the months of June, July, and August for the 2011 sampling season.

In the 2012 sampling season, the month of July as a sampling month was eliminated as this month did not add noteworthy data different from the months of June and August in 2011.

Vegetation Mapping

Patches were delineated on aerial photographs then converted to polygons using

ArcGIS software (ESRI 2011) to map patches across sites. Data compiled from the 2011 field season were used to cross-validate patch delineations of 2012 and adjusted as necessary.

Sample plots in each meadow from month to month and season to season were noted on aerial photographs, with overlaid delineated patches, using visual markers on the photograph and triangulation techniques to approximate position. GPS units were not used as the scale under investigation was too small to provide accurate plot positions. Positions noted on the aerial photograph patch maps were converted to points for each site in ArcGIS software.

Vegetation Characterization

Plants identified across the 2011 and 2012 field seasons were categorized by scientific name, Natural Resources Conservation Service (NRCS) species symbol (USDA

NRCS 2014), family status (USDA NRCS 2014), United States Fish and Wildlife Service

(USFWS) water affinity class (USFWS 1997), and plant type (Appendix A). Non-natives were also distinguished from invasive species (USDA NRCS 2014) in this study as they were found to have slightly different water affinities. In addition, for restoration it is important to differentiate which non-native species have the highest potential for spread so as to make the most efficient plan for future monitoring (SER 2004, Falk et al. 2006). USFWS wetland classifies species into one of six water affinity categories: obligate wetland (OBL), facultative wetland (FACW), facultative (FAC), facultative upland (FACU), obligate upland

(UPL), no agreement (NA), and no indicator (NI) (USFWS 1997). Specific descriptions and definitions for each of these water affinity types are in Appendix B. Plants were also classified by each species as annual (A) or perennial (P), though in some species there is a potential for both (A/P). In addition, plants were designated as forb (F), graminoid (G), or tree/shrub (T/S). There were two species that could not be identified by the end of the two field seasons, one a tree and the other a forb, and these species were excluded from all analyzed datasets.

After all possible species were identified, data were accumulated for each site’s patches over both field seasons (Appendix C). Patches were characterized by dominant species, dominant species vegetation type, average soil moisture category (see below), size, species richness, and average percent non-vegetated area. In this study, non-vegetated area was defined as any part of each plot that was not taken up by living vascular vegetation.

Thus, it could have been completely bare (with or without a cover of surface water), had some bryophyte growth, or had decaying duff from the previous growing season. The disturbed sites were observed to also have cobbles (64-256mm) taking up area in some of its plots. Generally non-vegetated areas in drier patches had harder packed soils than wetter patches, especially in the disturbed sites where topography was more extreme. Patches were designated by the dominant species vegetation types (>50% cover in a patch), categorized similarly as individual species designations above. However, not all vegetation types were observed and some patches were categorized as multi-species (MS). Multi species dominated patch types were designated if there was no discernible dominant species type taking up the majority of patch plant area. There were no patches dominated by annual/perennial plants or trees/shrubs. In summary there were five categories used: AG (annual graminoid), AF

(annual forb), PG (perennial graminoid), PF (perennial forb), and MS (multi-species).

Soil Moisture

The restoration at Big Trees State Park focused on raising the water table and restoring the hydrologic connections within the meadow. Comparing soil moisture of the restored meadow to the less-disturbed and disturbed meadows aided in evaluating the hydrologic connectivity and the restoration trajectory within the park. Soil moisture was estimated at each vegetation patch, within each meadow, across both sample months, using

16 the USDA hand method (USDA NRCS 2001) that categorizes soil moisture into five categories (0-25%, 25-50%, 50-75%, 75-100%, >100%). The hand method was applied to 2-

3 random spots in each patch depending on its size. In addition, one random soil sample was collected from each patch in tightly sealed bags then refrigerated in the lab until ready for further analysis. Soil samples of approximately 118 cm3 were taken at 10cm below the humus horizon, taking the bottom 6-7cm and replacing the top 3-4cm so as to minimize disturbance to the seed bank. Subsequently the oven drying method (Gardner 1986) for estimating soil moisture was used to verify hand method estimations. The oven drying procedures were to first remove soil from bags, making sure any condensate inside the bag was reabsorbed into the soil by shaking the soil around in the bag, then to weigh each soil sample as quickly as possible. Soil samples were then baked for 24hrs at 104°C, then reweighed to estimate moisture loss. The soil moisture estimations were done during the

2012 field season and thus all data organized by moisture conditions only reflect this particular sampling year. In addition, soil moisture categories were overlain onto patch delineation maps to investigate hydrological connections to vegetation patterns. Moisture maps were created based on average moisture between June and August moisture estimates.

In addition, June moisture maps were inlaid to also get an idea of the moisture pattern wettest sampling month. This wettest month is important as it sets the stage for phenology and pattern of vegetation for the season.

Data Analysis and Statistics

Data Preparation

Percent cover for all species present in each quadrat sampled through the 2011-

2012 sampling seasons was recorded along with the associated sampling month, patch, and

17 meadow. For each sampling year, quadrats were sorted by patch for each meadow, and each species percent cover present in those quadrats was averaged to represent the average percent cover of each species for the entire growing season (June and August) for that particular patch. Delineated patch polygons were used to calculate each patch area across all sites. Area for each present species in a patch was estimated by multiplying its average percent cover value by patch area. These species area values were then divided by the total area of the study meadow to get a relative area for that species. Both the species estimated area of cover and relative area were used as a basis for answering either meadow level inferences (where values would be grouped together by site, by native status, by plant type, or by USGS water affinity) or patch level inferences (where values would be used individually or grouped by soil moisture categories). It is important to note that investigations of patterns based on hydrology only used 2012 data for comparisons as that was the only year hydrologic data was collected. Unless otherwise stated, most other investigated variables utilized averaged species estimated area of cover and relative area between the 2011 and 2012 sampling seasons.

Richness Comparisons

Whenever data were grouped, as described above, and the same species were present in the groups being combined, the values for their areas were summed (For example, if Collomia linearis was present in Patch 1 and 5, then its total estimated area for each patch would be summed to get one number). In such cases, species relative areas also had to be recalculated by taking the summation of area in each group of interest and dividing the total site area. Other relative areas within groups were calculated by taking the sum of the area for the group of interest and divided by the total possible area for that group. For native status,

18 plant type, USGS water affinities, and soil moisture categories, summations of areas were then used to calculate species richness in each group using a presence/absence counting command. Note that all of the calculations were first based on estimations of average percent cover values for each species in a patch. Thus, instances where relative areas are presented within categories, these areas would not add up to 100%. Data analysis was done this way to reflect Sierra Nevada meadow natural history where there is within patch uniformity (follows the definition that patches are visually distinct and repeated entities described in vegetation sampling section above), and between patch heterogeneity where covers of species are much more variable.

A chi-square test of independence was run to compare observed richness values between native and exotic (summed invasive and non-native species) across the sites. It was hypothesized that sites and status richness would be associated due to disturbed hydrology potentially interrupting montane meadow characteristic of low exotic to native ratio.

Assumptions for chi-square test of independence were met based on having nominal data with multiple independent groups. Raw observations were entered into R-studio (RStudio

2015) for analysis.

Precipitation Comparisons

As sites were split between two watersheds and two sampling years, individual yearly precipitation totals for each watershed’s weather station were taken from the National

Oceanic and Atmospheric Administration (NOAA). Yearly estimates were compared to the average total precipitation data compiled from each watersheds gauging station since their operation. NOAA changed how they reported total precipitation in 2001 from a calendar year report to a water year (WY) report (October-September) (NOAA 2012). As such, this system

19 takes into account the precipitation that falls as snow during autumn and winter but does not drain until the following spring and summer. In sampling this way there is more confidence that totals for each water year are independent, this is especially important for statistical comparisons. As the WY system started in 2002, there was only nine sampling years up until the start of this research. Using WY total precipitation data, averages and standard deviations were calculated for each watershed. A paired t-test was conducted in R-studio (RStudio

2015) to investigate differences in precipitation between watersheds. One sample t-tests were also computed in R-studio to compare precipitation differences for each sample year to the running precipitation averages (within a watershed for the WY system) to get an idea of how wet or dry the sample year was.

Non-vegetated Comparisons

Abiotic and biotic differences between sample seasons also lead to an investigation of percent non-vegetated cover variation across sites and sample years. Percent non-vegetated cover was summed for every plot by subtracting the sum of plant percent covers in that plot and subtracting it from 100% as there were no instances where cover was over 100%. All of these raw percent non-vegetated cover values were then used as the response in a longitudinal model analysis in R-studio (RStudio 2015) to test how year

(repeated measure) and site influenced non-vegetated area patterns. It was hypothesized that year would significantly affect non-vegetated area at the sites due to observed differences of total precipitation between sample years. Assumptions for longitudinal model analysis are met based on quantitative response data, year covariate, and sites assumed being independent.

For each sampling year, non-vegetated percent cover for each plot was averaged across all plots sampled in a patch to represent the mean percent non-vegetated cover for that patch for that particular year. These means were then multiplied by patch areas to get the estimated area of non-vegetated cover per patch, and then divided by the area of the meadow to get relative cover values (same calculations used for percent cover values for each species described above). These area cover values and relative cover values were used to answer meadow level inferences or patch level inferences as described above.

CHAPTER III

RESULTS

Variation Between Watersheds and Between Sampling Years

Apparent differences in moisture patterns across sites in different watersheds prompted an investigation of total water year precipitation differences (Table 2). The Upper

Calaveras had significantly greater average annual precipitation than the Upper Merced (t =

7.46, df = 10, p <0.001). When comparing individual sampling year precipitation for each watershed to the overall average annual precipitation, 2011 was a significant wet year (Upper

Calaveras: t = -5.43, df = 8, p < 0.001; Upper Merced: t = -6.50, df = 8, p <0.001) and 2012 was a significant drier year (Upper Calaveras: t = 3.66, df = 8, p = 0.003; Upper Merced: t =

3.87, df = 8, p = 0.002).

Table 2. Total precipitation averages (inches), using the current water year system, between watersheds for both sampling years.

Upper Calaveras Upper Merced Average *51.1 33.90 StDev 14.24 9.74 n (years) 9.00 9.00 2011 **76.9 **55.0 2012 *33.7 *21.4 *Indicates a significant higher value than the Upper Merced. **Represent statistical testing between the WY watershed averages to each individual year. The quantity of asterisks indicates the level of significance (**P-value<0.001, *P- value<0.01).

This variation in precipitation between years was evident not only because of noted moisture differences but also in observed plant phenology and biomass variations

22 between sampling years. For example, some species (esp. Solidago Canadensis – Goldenrod) could not be identified the first sampling season, as the species had not even begun flowering by August (last sampling month). In contrast, the following sampling season (2012) most of these species were already in flower and by June most had already gone to seed. Another observed difference was overall increased average sizes (biomass) of plants observed through the sampling season. For example, Lupinus latifolius var. columbianus (lupin) and Mimulus guttatus (common monkeyflower) had estimated growth above their maximum height ranges in 2011 while in 2012 their heights were much more average. It is important to note that all of these observations were not officially measured but rather photographed and documented in a field notebook. Thus, no official data was taken to further scrutinize these differences.

Vegetation and Soil Moisture

The total number of patches within a site corresponded with the overall size of each site. Thus, the restored meadow (BT), the largest site in terms of area, also had the most patches, followed by the two disturbed meadows (DR, HF) (Table 3). The less-disturbed sites (EL, HD) were the smallest meadows with less than half the area as the restored site and also the fewest patches. Overall, the disturbed and restored sites had similar counts of patch types though they had notably more than the less disturbed sites.

Patches were further categorized by soil moisture categories based on soil moisture estimations from soil samples in 2012 (Table 4, Figure 6). In general, the restored site had more patch area in higher moisture categories followed by the disturbed sites, and lastly the less-disturbed sites had the driest patch areas. The disturbed sites (DR and HF) had the highest patch area and patch count in the 0-25% and 25-50% moisture categories, with approximately half of their total area in the driest soil moisture category. The majority of

Table 3. Total estimated meadow area (m2) and number of patches and (vegetation patch types) surveyed for 2012. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed.

Patch Count Site Area (Types) DR 7416.0 28 (13) HF 10635.9 35 (15) BT 20170.8 43 (14) EL 7120.6 13 (9) HD 2951.7 18 (9)

vegetation patch types for the disturbed sites were also in the three driest categories. For example, for HF there were 15 different patches in the 0-25% category and that represented nine different vegetation types. In contrast, more than half of the area of the restored BT was in the 25-50% moisture category and approaching 20% of the area was in each of the 50-75% and 75-100% moisture categories. BT also had the highest number of patches in both the 25-

50% and 75-100% moisture categories with 14 patches classified as four vegetation patch types patches in the 25-50% category taking over half of its total area. Though overall a smaller area (18%), there were also 15 patches classified among four vegetation patch types in the 75-100% moisture category. The less disturbed sites (EL and HD) had the highest numbers of patches, patch types, and areas in the 0-25% moisture category with over 90% of their total area in the driest category.

Variation in moisture patterns across the study meadows can also be detected in the 2012 moisture maps (Figure 7a-e). DR (Figure 7a.) and HF (Figure 7b.) were identified as hydrologically disturbed as evidenced by the somewhat restricted waterways running through them. Compared to the disturbed sites, the restored site (Figure 7c.) had more consistent

Table 4. Study sites classified by 2012 patch soil moisture categories. Within each moisture category, the area (m2) and (relative area) as well as the number of patches and (the number of different patch types) are listed. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed.

0-25% 25-50% 50-75% 75-100% >100% Patch Patch Patch Patch Area Count Area Count Area Count Area Count Area Patch Count Site (%Area) (Types) (%Area) (Types) (%Area) (Types) (%Area) (Types) (%Area) (Types) DR 3549.9 (47.9) 9 (6) 2216.2 (29.9) 6 (5) 662.7 (8.9) 8 (5) 871.9 (11.7) 4 (2) 115.2 (1.5) 1 (1) HF 5627.8 (52.9) 15 (9) 2314.8 (21.8) 7 (4) 283.8 (2.7) 5 (5) 1276.4 (12.0) 7 (4) 1133.0 (10.6) 2 (2) BT 764.3 (3.8) 4 (2) 11660.9 (58.3) 14 (4) 3853.3 (19.3) 8 (6) 3716.5 (18.4) 15 (4) 175.7 (<1) 2 (2) EL 6429.6 (90.3) 11 (8) 691.0 (9.7) 2 (2) 0.0 (0.0) 0 (0) 0.0 (0.0) 0 (0) 0.0 (0.0) 0 (0) HD 2735.2 (92.7) 13 (8) 167.5 (5.7) 2 (1) 49.0 (1.7) 3 (1) 0.0 (0.0) 0 (0) 0.0 (0.0) 0 (0)

Figure 6. Total relative area of patches classified by 2012 soil moisture categories for each site. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed.

0-25 25-50 50-75 75-100 >100

100 90 80

70 60 50 40 Relative Relative area 30 20 10 0 DR HF BT EL HD Site

moisture connectivity, hence less extreme moisture differences between adjacent patches throughout the site. In contrast, the two less disturbed sites ((EL, Figure 7d) and (HD, Figure

7e)) were consistently drier and had less variation in moisture patterns. From the maps and moisture levels (Table 4), it is evident that BT had more patches with higher moisture values compared to both the disturbed and less-disturbed sites.

Vegetation maps of the sites (Figure 8a-e.) illustrate the different vegetation types that dominated each patch averaged across both sampling seasons. The dominate species, number of patches dominated by each species-type, and the area they encompassed was also summarized for each site (Appendix D). Disturbed sites had the greatest number of patches in perennial forbs (PF) and perennial graminoids (PG) categories but the majority of the site area was distributed between PF, PG and mixed-species (MS) dominated patches. The restored site also had the most patches in PF and PG dominated patches but PG dominated

26 patches took up the majority of its area. The less-disturbed sites had the majority of their area taken up by MS dominated patches. For EL there was no plant type that dominated in terms of number of patches but for HD the majority of patches were dominated by PG type vegetation.

When comparing both the vegetation patch (Figure 8a-e.) and moisture maps

(Figure 7a-e.) there is a pattern that moist patches tended to be single vegetation type dominated while drier patch types tended to be multi-species dominated patch types. This trend is evident most in the disturbed sites DR and HF. For example, there is a clear overlap of the higher moisture patches and single species dominated (unfilled or solid shapes) patches while the less moist were classified as MS (blank) patches. Also for HF the more moisture patches were also single vegetation type dominated while the drier patches, particularly in the center, were MS patches. The restored site however, does not uphold this pattern and of the two small MS patches it has, they are both classified with the highest moisture values. The less-disturbed sites have similar patterns to the disturbed in that the MS patches they have all come from the lowest moisture patch types. For these meadows, it was not always the case that the heaviest moisture patches were single vegetation type dominated, however it was still a strong predictor of moisture influencing vegetation type.

Richness

A total of 123 species and 41 families were identified across all sites through the duration of this research (Table 5, Appendix A). Though they were not the largest, the two disturbed sites, had the highest total species and family richness followed by the restored site, then the less-disturbed sites with the lowest richness. HD had an especially low richness

Figure 7a. DR.

Figures 7a-e: Average soil moisture maps across the sampling season, with inset for June (wettest sampling month) moisture, for the delineated vegetation patches based on field sampling in June 2012 at each site: (a.) DR, (b.) HF, (c.) BT, (d.) EL, (e.) HD. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. Note differences in map scales, 27 especially between the sites in different watersheds.

Figure 7b. HF.

Figure 7c. BT.

Figure 7d. EL.

Figure 7e. HD.

Figure 8a. DR.

Figure 8a-e. Dominant plant type, or not, across both sample seasons at (a.) DR, (b.) HF, (c.) BT, (d.) EL, and (e.) HD. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. Note differences in map scales, especially 32 between the sites in different watersheds.

Figure 8b. HF.

Figure 8c. BT.

Figure 8d. EL.

Figure 8e. HD. 36

37 compared to the other sites with approximately half the richness compared to the EL, the other less disturbed site. However, a chi-square test of independence comparing native and exotic (summed non-native and invasive totals from Table 5) species richness found no associations with sites (X2 = 1.07, df = 4, p = 0.89). Despite these similar patterns of richness and species status across the sites, it was apparent that native species richness was notably higher across all meadows. Non-native percent total richness values were similar across sites, with EL having slightly higher percentage than the rest. Invasive percent total richness values showed slightly larger percentages with increasing disturbance, though HD was the only site with a more noteworthy (lower) value.

Table 5. Richness and (percent total richness) for native, invasive, and non-native species as well as total species richness and family richness across both sampling seasons. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed.

Site Native Non-native Invasive Total Family DR 53 (74.6) 6 (8.5) 12 (16.9) 71 28 HF 43 (71.7) 6 (10.0) 11 (18.3) 60 25 BT 39 (75.0) 5 (9.6) 8 (15.4) 52 21 EL 37 (74.0) 6 (12.0) 7 (14.0) 50 16 HD 19 (82.6) 2 (8.7) 2 (8.7) 23 11 Total 98 (79.7) 12 (9.7) 13 (10.6) 123 41

As data were collected between notably different water years (Table 2), a comparison was also made between cumulative species richness between and across both sampling years and sites (Table 6). The first sampling year noted increased richness values with increased disturbance of the sites. However, in the second sampling year the disturbed

and restored sites were similarly large, but still larger than the less-disturbed sites. There was 37

an observed decrease in richness from 2011 to 2012 for all sites.

Table 6. Species richness for each site between sampling years (2011 and 2012). The site HD has no measurements for 2011 as the site was switched in 2012 to a new location. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed.

Site 2011 2012 DR 59 44 HF 52 44 BT 45 43 EL 35 33 HD -- 25

For 2012, species richness were also calculated across patch soil moisture categories for each site (Table 7). These values are presented along with percent area estimations (Table 3) to allow comparisons of area to richness. The disturbed sites had the largest areas and highest richness values in the 0-25% moisture category. The restored sites

25-50% category had the majority of the total area but had a similar richness value compared to the two moisture categories above it. The restored sites driest and wettest moisture categories had the smallest area and richness values. The less-disturbed sites had a similar trend to the disturbed sites where the 0-25% moisture category was associated with the highest area and richness values (Table 7).

For each meadow, species richness within the different plant-type categories

(growth forms) was also summed along with the average percent of total area encompassed within each category between sampling years (Table 8). For example, in 2011 for DR there were four species in the AG category, which comprised 2.2% of the total average area in that site. Across the sites the most numerous species vegetation types, which also had the average

percent of total areas were PG and PF. However, the disturbed meadows also had high 38

average percent of total area for AF despite not having many species in this category.

Table 7. Percent of meadow area in each soil moisture category and the accompanying species richness in 2012 for each site. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. Note the zero values for the higher moisture categories for EL and HD reflect the lack of patches with those moisture designations within the site.

0-25% 25-50% 50-75% 75-100% >100% Site % Area Richness % Area Richness % Area Richness % Area Richness % Area Richness DR 47.9 50 29.9 46 8.9 38 11.7 36 1.5 17 HF 52.9 52 21.8 34 2.7 23 12.0 37 10.6 12 BT 3.8 22 58.3 37 19.3 38 18.4 37 <1.0 23 EL 90.3 46 9.7 20 0.0 0.0 0.0 0.0 0.0 0.0 HD 92.7 24 5.7 6 1.7 2 0.0 0.0 0.0 0.0

Table 8. Species richness and (average percent of total area) of plant types across sampling years (AG = annual graminoid, AF = annual forb, AF/PF = annual/perennial forb, PG = perennial graminoid, PF = perennial forb, T/S = tree/shrub) in each site. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. The relative percentages don’t total 100 because non-vegetated percentages are missing and the way areas and relative areas were estimated (outlined in the methods).

AG AF AF/PF PG PF T/S Site 2011 2012 2011 2012 2011 2012 2011 2012 2011 2012 2011 2012 DR 4 (2.20) 2 (1.96) 6 (21.48) 3 (8.19) 1 (0.09) 1 (0.23) 16 (23.14) 15 (26.53) 31 (20.21) 22 (12.06) 1 (0.03) 1 (0.14) HF 4 (1.77) 1 (1.28) 7 (25.36) 6 (9.71) 0 (0.00) 0 (0.00) 16 (13.91) 17 (17.82) 24 (27.99) 20 (25.04) 1 (0.03) 0 (0.00) BT 3 (3.73) 2 (5.94) 9 (3.46) 5 (1.48) 1 (0.55) 1 (0.04) 12 (14.66) 15 (13.30) 19 (36.86) 20 (26.70) 1 (0.01) 0 (0.00) EL 5 (5.22) 3 (8.78) 4 (1.14) 3 (0.20) 0 (0.00) 1 (0.01) 10 (30.31) 12 (13.07) 16 (19.46) 13 (8.99) 0 (0.00) 1 (0.01) HD -- (--) 2 (0.06) -- (--) 2 (0.06) -- (--) 0 (0.00) -- (--) 9 (31.48) -- (--) 11 (11.14) -- (--) 1 (0.07)

Between years there were observed decreases in richness and the average percent of total area amongst the plant-type categories.

To evaluate the relationship between plant growth forms and moisture, the percent area of each plant type in each moisture category was calculated for each site (Table 9,

Appendix Ea.). For all the sites PG and PF occupied the greatest percentage of area across the moisture categories. The restored site had consistently higher percentage in the PF plant type compared to PG for all moisture categories except the wettest category. Though relatively low in percentage, BT also had consistently more AG present across its moisture categories compared to the other sites.

For each sampling year plant species were grouped based on their USGS wetland classification, the total richness, and average percent total area was determined in each category (Table 10). Across the sites the OBL category was the most rich and had the highest average percentages of total areas followed by FAC. Although for the disturbed sites FACW species also had strong values. FACW also had high richness values in the restored site but had an overall smaller relative area. The NI category of wetness was mixed in terms of relations of richness to average percent of total area across sites, though there were noteworthy contributions from this category for all the sites. Between years there were observed decreases in richness and the average percent of total area amongst the wetland classification categories.

To get a clearer representation of the extent of species wetness affinities associated with site moisture patterns, the average percent total area occupied by plants in specific wetland categories within each moisture patch was estimated across patch soil moisture categories (Table 11, Appendix Eb). In general there were no consistent trends

Table 9. The average percentage of total area in each moisture category across occupied by a particular plant type in 2012 for each site. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. The relative percentages do not total 100% because of the way plant areas and relative areas were estimated using averages across sample months and multipliers (outlined in the methods). EX: There was a total of 3.2% AG species found on average across sample months in the 0-25% moisture category of DR in 2012.

Moisture Plant Type Site category AG AF AF/PF PG PF T/S 0-25% 3.2 11.5 <1 26.4 8.2 <1 25-50% 1.1 4.5 0.0 35.0 13.8 0.0 DR 50-75% 0.0 4.3 1.3 18.3 4.1 0.0 75-100% 0.7 7.0 <1 14.1 25.6 0.0 >100% 1.0 9.0 0.0 8.0 42.0 0.0 0-25% 2.2 10.8 0.0 14.0 19.3 0.0 25-50% <1 11.4 0.0 18.2 42.2 0.0 HF 50-75% <1 <1 0.0 17.2 13.5 0.0 75-100% <1 7.4 0.0 31.9 15.4 0.0 >100% <1 5.6 0.0 20.2 32.4 0.0 0-25% 2.8 <1 0.0 13.5 38.9 0.0 25-50% 8.3 <1 0.0 12.4 23.6 0.0 BT 50-75% 3.3 4.5 0.0 11.4 34.2 0.0 75-100% 2.0 <1 0.0 16.4 25.5 0.0 >100% <1 1.2 4.1 37.7 12.2 0.0 0-25% 8.9 <1 <1 13.4 7.4 <1 EL 25-50% 7.7 <1 0.0 10.3 24.0 0.0 0-25% 21.8 <1 0.0 27.2 12.0 <1 HD 25-50% 0.0 0.0 0.0 85.3 <1 0.0 50-75% 0.0 0.0 0.0 84.7 0.0 0.0

across sites. At the disturbed sites, OBL tended to dominate the average percent total areas across all moisture categories. However patterns were different between the disturbed sites with DR having higher densities in lower moisture soils, and HF having the higher densities in the middle and highest moisture categories. In the restored site relative area covered by

Table 10. Species richness and (average percent of total area) categorized according to the USGS wetland classification categories across sampling years (OBL= obligate, FACW= facultative wetland, FAC= facultative, FACU= facultative upland, UPL= upland, NI= no indicator). Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed.

OBL FACW FAC FACU UPL NI Site 2011 2012 2011 2012 2011 2012 2011 2012 2011 2012 2011 2012 DR 16 (46.23) 13 (32.19) 13 (7.87) 11 (4.15) 11 (4.61) 5 (2.15) 5 (4.25) 4 (3.74) 3 (1.15) 2 (0.07) 11 (3.04) 9 (6.82) HF 12 (36.28) 12 (16.57) 9 (7.74) 8 (7.23) 15 (18.22) 12 (17.23) 6 (3.77) 5 (5.57) 1 (1.29) 1 (0.12) 9 (1.77) 6 (7.13) BT 12 (26.66) 10 (8.95) 8 (4.18) 8 (5.19) 10 (12.51) 11 (8.88) 6 (10.44) 6 (10.79) 3 (0.38) 2 (0.31) 6 (5.10) 6 (13.27) EL 6 (22.88) 7 (8.82) 4 (1.60) 7 (4.11) 10 (16.28) 7 (5.43) 6 (6.96) 2 (0.88) 1 (0.07) 2 (0.06) 9 (8.45) 8 (11.77) HD -- (--) 3 (14.26) -- (--) 7 (4.83) -- (--) 7 (36.52) -- (--) 4 (5.94) -- (--) 1 (0.05) -- (--) 3 (1.39) Note. HD was not sampled in 2011.

Table 11. Average percent of total area within each site’s 2012 patch moisture categories occupied by plant species classified by the USGS wetland classification system. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. The relative percentages do not total 100% because of the way plant areas and relative areas were estimated using averages across sample months and multipliers (outlined in the methods). EX: There was a total of 34.7% OBL species found on average across sample months in the 0-25% moisture category of DR in 2012.

Moisture Wetland Classifications Site category OBL FACW FAC FACU UPL NI 0-25% 34.7 <1 2.6 3.9 <1 7.6 25-50% 34.3 7.5 2.3 4.2 <1 6.0 DR 50-75% 21.8 3.8 <1 1.1 0.0 1.0 75-100% 27.3 6.2 1.3 2.9 0.0 10.2 >100% 11.0 30.0 3.0 10.0 0.0 6.0 0-25% 12.5 4.7 11.0 8.9 <1 9.0 25-50% 15.4 13.5 37.2 3.6 0.0 2.5 HF 50-75% 26.9 0.0 1.7 <1 0.0 2.8 75-100% 14.9 10.4 15.4 <1 0.0 13.7 >100% 38.5 5.4 13.6 <1 0.0 1.0 0-25% 1.5 4.9 20.9 24.5 0.0 3.8 25-50% 2.1 4.6 10.3 11.4 <1 16.4 BT 50-75% 17.5 9.4 5.3 8.3 <1 12.2 75-100% 21.3 2.2 5.6 8.5 <1 6.6 >100% 36.9 11.0 1.7 5.0 <1 <1 0-25% 7.0 4.0 5.7 1.0 <1 12.1 EL 25-50% 25.9 4.8 2.9 <1 0.0 8.2 0-25% 15.3 5.2 32.8 6.4 <1 1.5 HD 25-50% <1 <1 84.7 0.0 0.0 0.0 50-75% 1.4 0.0 68.9 0.0 0.0 0.0

FAC and FACU species decreased with increasing moisture; however NI had higher densities in the second and third moisture category. In the three wettest categories, BT had the greatest relative area occupied by OBL species. Less-disturbed EL, which only had patches in the two lower moisture categories, had higher relative areas of OBL species in the wetter of the two categories.

Given lower average percent total areas in Table 11, I further investigated patterns in average percentage of non-vegetated area across sites and between years (Table 12).

Between 2011 and 2012 across all sites, the percentage of non-vegetated ground increased on average by more than 20% with BT having the smallest percentage increase of 12%. Within years, in 2011, the less-disturbed site EL had the highest percentage of non-vegetated area followed by the restored site. The disturbed sites had the lowest relative non-vegetated areas.

However, overall the greatest area of actual non-vegetated ground was at BT followed by DR and HF. A longitudinal model analysis, with the repeated measure of sample year, was run to compare sites to the average total percent non-vegetated values (as described in methods).

Between the two sample years it discovered significant differences between sites total percent non-vegetated values (F = 13.77, df = 3, p < 0.001).

The average total percentage non-vegetated area at a site in each soil moisture category and within each moisture category was also estimated for the 2012 (Figure 9,

Appendix F). All sites had the largest total non-vegetated areas in the moisture category with the highest total area. Thus, all sites had the most non-vegetated area in the driest moisture categories apart from BT, which had the highest area in 25-50% moisture category. Similar trends were also reflected for the largest average total percentage non-vegetated areas across the sites, excluding DR that had its highest average total percentage non-vegetated area in the

75-100% moisture category where, on average, over 50% of the area was non-vegetated. For all sites, the lowest average total percentage area of non-vegetated ground within moisture categories were found in the middle moisture category.

Table 12. The average area of non-vegetated ground (m2) and (average percentage of total area) at site scale between years. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed.

Site 2011 2012 DR 2149.0 (29.0) 3400.6 (45.8) HF 1199.2 (11.3) 4580.3 (43.1) BT 7285.0 (36.4) 9645.4 (48.3) EL 3330.5 (46.8) 4901.0 (68.8) HD -- (--) 1143.5 (38.7)

Invasive and Non-Native Species

Total area and relative area occupied by invasive, non-native, and native species were summarized for both sampling years at each site (Table 13). Across sites and between years, less than a tenth of the total area of each site was covered with invasive and non-native species. Natives in comparison, hold higher areas for both sampling years in all sites but EL.

In 2011, the wetter of the two years of the study, the less-disturbed site EL had the highest invasive and non-native areas and the lowest native area. The restored site had the second largest invasive and non-native areas and the second lowest native areas. Surprisingly, the disturbed sites had the least overall area of invasive and non-native species and the highest areas of native species. The following sampling year showed the same general trend across sites, though the total areas were less than they were in 2011. HD also followed the trends for invasive and non-native species however it native area was the largest of any other meadow.

It is important to remember that for HD, 2012 was the only year of sampling and thus trends between years cannot be determined. The HF non-native area was the only category not to decrease from the 2011 to the 2012 sampling season. In general, there were higher overall relative areas for invasive compared to non-native species across the sites aside from HD’s

2012 values.

0-25 25-50 50-75 75-100 >100

50 VegetatedTotal Area - 40

Average Average PercentNon 10

0 DR HF BT EL HD SIte

Figure 9. Average percent non-vegetated total area across moisture categories at each site for 2012. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed.

As previously noted, at the meadow scale cumulative richness showed similar percentages of invasive and non-native species across all sites (Table 5). To better understand the community association of the invasive and non-native species, they were categorized by the vegetation types (Table 14) and USDA wetland classification categories (Table 15).

Overall the PF vegetation association had the greatest number of invasive and non-native species and also comprised the greatest percentage of species richness at each site. Invasive species were primarily associated with PG and PF community types (Table 14). Most sites,

Table 13. Average area (m2) and (average percent of total area) occupied by invasive, non-native, and native species for each sampling year at each site. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed.

Invasive Non-native Native Site 2011 2012 2011 2012 2011 2012 DR 112.01 (1.51) 32.60 (0.44) 63.72 (0.86) 6.56 (0.09) 1896.38 (25.57) 1527.06 (20.59) HF 283.16 (2.66) 182.23 (1.71) 21.26 (0.20) 54.06 (0.51) 2499.96 (23.51) 2211.36 (20.79) BT 1296.25 (6.48) 175.28 (0.88) 162.46 (0.81) 93.46 (0.47) 2638.60 (13.20) 2019.46 (10.10) EL 721.21 (10.13) 178.31 (2.50) 111.41 (1.56) 38.76 (0.54) 613.59 (8.62) 384.94 (5.41) HD -- (--) 182.58 (6.19) -- (--) 256.75 (8.70) -- (--) 681.76 (23.10)

Table 14. The number of invasive and non-native species by plant type and (percent of total species richness for each plant type) at each site. In addition, total richness values for each plant type was summed across all sites. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. See vegetative sampling section in method for abbreviations.

AG AF AF/PF PG PF T/S Site I NN I NN I NN I NN I NN I NN DR 1 (1.4) 1 (1.4) 1 (1.4) 3 (4.2) 0 (0.0) 0 (0.0) 3 (4.2) 1 (1.4) 7 (9.9) 1 (4.5) 0 (0.0) 0 (0.0) HF 1 (1.7) 1 (1.7) 1 (1.7) 2 (3.3) 0 (0.0) 0 (0.0) 3 (5.0) 1 (1.7) 6 (10.0) 1 (1.7) 0 (0.0) 1 (1.7) BT 1 (1.9) 0 (0.0) 0 (0.0) 2 (3.8) 0 (0.0) 0 (0.0) 4 (7.7) 1 (1.9) 3 (5.8) 2 (3.8) 0 (0.0) 0 (0.0) EL 1 (2.0) 1 (2.0) 0 (0.0) 2 (4.0) 0 (0.0) 0 (0.0) 3 (6.0) 0 (0.0) 3 (6.0) 3 (6.0) 0 (0.0) 0 (0.0) HD 0 (0.0) 1 (4.3) 0 (0.0) 0 (0.0) 0 (0.0) 0 (0.0) 1 (4.3) 1 (4.3) 1 (4.3) 0 (0.0) 0 (0.0) 0 (0.0) Total 1 1 1 5 0 0 4 1 7 4 0 1

Table 15. Invasive and non-native species counts and (percent of total species richness at each site in the USDA wetland classification categories) across both sampling seasons. In addition, total richness values for each USDA wetland classification category was summed across all sites. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. See vegetative sampling section in method for abbreviations.

OBL FACW FAC FACU UPL NI Site I NN I NN I NN I NN I NN I NN DR 0 (0.0) 1 (1.4) 1 (1.4) 0 (0.0) 6 (8.5) 3 (4.2) 3 (4.2) 1 (1.4) 0 (0.0) 0 (0.0) 2 (2.8) 1 (1.4) HF 0 (0.0) 1 (1.7) 1 (1.7) 0 (0.0) 6 (10.0) 2 (3.3) 3 (5.0) 1 (1.7) 0 (0.0) 0 (0.0) 1 (1.7) 2 (3.3) BT 0 (0.0) 1 (1.9) 0 (0.0) 0 (0.0) 4 (7.7) 2 (3.8) 4 (7.7) 1 (1.9) 0 (0.0) 1 (1.9) 0 (0.0) 0 (0.0) EL 0 (0.0) 1 (2.0) 1 (2.0) 0 (0.0) 4 (8.0) 1 (2.0) 2 (4.0) 1 (2.0) 0 (0.0) 1 (2.0) 0 (0.0) 2 (4.0) HD 0 (0.0) 0 (0.0) 0 (0.0) 0 (0.0) 1 (4.3) 1 (4.3) 1 (4.3) 0 (0.0) 0 (0.0) 0 (0.0) 0 (0.0) 1 (4.3) Total 0 2 1 0 6 4 4 2 0 1 2 3

51 apart from HD, had one AG invasive species but there were no AF/PF or T/S invasive species in any meadow. For non-native species there was more variation in plant type associations compared to invasive species. Overall, non-native species were more often associated with AF and PF communities but most sites also had AG and PG non-native species present though the overall percentage compared to total species richness for each site was relatively low.

In the USGS wetland classification categories, the greatest number and the largest percentage of invasive species across all sites were found to be in the FAC and FACU categories (Table 15). Notably, there were no invasive species present in any site in the OBL or UPL category. The only wetland category that had non-native species present in every site was FAC. Though low in numbers, OBL and FACU non-native species were present at all sites except HD, and NI species were present at all sites but the restored. Only DR and BT had an UPL non-native species, and there were no FACW category non-native species present at any sites. In general, invasive species were mostly FAC and FACU species with no

UPL species types, and non-natives were mainly FAC with no FACW species types.

For each invasive and non-native species (organized by wetland classification), the average area, and average percent total area, were also calculated across each meadow for each sampling year. However, most values covered by these species related to average percent total area cover were less than one percent. To examine patterns related to the more common invasive and non-native species between years, only species with at least one value over a one percent average total cover are presented in Table 16. Of the twenty-five invasive and non-native species identified, only four invasive species (PHAQ (Phalaris aquatica),

ANOD (Anthoxanthum odoratum), RUCA3 (Rumex acetosella), HOLA (Holcus lanatus))

Table 16. Average area (m2) and (average percent of total area (%)) across both sampling seasons occupied by the most prevalent invasive and non-native species for each meadow organized by their wetness affinities. Invasive and non-native species listed had at least one value where their percent cover is over 1%. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed.

FAC FACU Yr Site PHAQ RUCA3 HOLA POCO MOVE ANOD DR 29.8 (<1) 57.5 (<1) 14.1 (<1) 0.0 (0.0) 0.0 (0.0) 0.0 (0.0)

HF 77.4 (<1) 61.4 (<1) 43.8 (<1) 0.0 (0.0) 0.0 (0.0) 0.0 (0.0)

BT 937.0 (4.7) 2.9 (<1) 4.1 (<1) 0.0 (0.0) 42.0 (<1) 296.2 (1.5) 2011 EL 478.3 (6.7) 177.7 (2.5) 5.2 (<1) 0.0 (0.0) 73.7 (1.0) 1.5 (<1) HD -- (--) -- (--) -- (--) -- (--) -- (--) -- (--) DR 0.0 (0.0) 0.0 (0.0) 0.0 (0.0) 0.0 (0.0) 0.0 (0.0) 32.6 (<1)

HF 0.0 (0.0) 89.2 (<1) 37.3 (<1) 7.0 (<1) 0.0 (0.0) 0.20 (<1)

BT 30.3 (<1) 11.2 (<1) 0.00 (0.00) 6.2 (<1) 23.3 (<1) 120.2 (<1) 2012 EL 0.0 (0.0) 160.8 (2.2) 13.4 (<1) 0.0 (0.0) 0.7 (<1) 0.0 (0.0) HD 0.0 (0.0) 0.0 (0.0) 179.4 (6.1) 239.7 (8.1) 0.0 (0.0) 0.0 (0.0) Key: PHAQ (Phalaris aquatica), ANOD (Anthoxanthum odoratum), RUCA3 (Rumex acetosella), HOLA (Holcus lanatus), POCO (Poa compressa) and MOVE (Mollugo verticilla)

and two non-native species (POCO (Poa compressa) and MOVE (Mollugo verticilla)) had areas that represented an average over one percent total area of a site. It is notable that of the most common invasive and non-native species noted in Table 15 that five of six are FAC species.

With invasive species, the disturbed sites had no invasive species that had average values over one percent in either sampling year while the less-disturbed site EL had the highest values followed by the restored site. For non-native species, MOVE had the only value over one percent in 2011, in EL and POCO had the only value over one percent in

2012 in HD. The most common species found between sites and years were: PHAQ,

RUCA3, and HOLA. Between years, areas of invasive and non-native species decreased or stayed roughly the same areas across sites.

Moving to site patch scale, the total invasive and non-native average areas, and average percent total areas, were grouped by patch soil moisture categories (Table 17a-b). In general, invasive and non-native species occupied a relatively small area for any of the individual moisture categories across the study sites. For invasive species, sites had the highest average percent total area of invasive species in the driest category excepting HD, which had extremely high values in the second and third soil moisture categories. This being said, these two moisture categories took up less than 8% of the average total area for the site.

The restored site had the next highest value of invasive species across the sites but again the relative areas were quite small. For non-native species there was less overall area and densities than invasive species. The only site to have notable densities of non-native species was HD, and these densities were found in the driest soil moisture category.

Table 17a-b. Relative area for each of the moisture categories in each site, total average area (m2) and the (average percent of total area for each moisture patch occupied) by invasive (a) and non-native (b) species across the 2012 sampling season. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. a. 0-25% 25-50% 50-75% 75-100% >100% Area (m2) % of (% of Patch Site site Occupied) % Area Area (%) % Area Area (%) % Area Area (%) % Area Area (%) DR 47.9 25.0 (<1) 29.9 0.0 (0.0) 8.9 0.6 (<1) 11.7 7.0 (<1) 1.5 0.0 (0.0) HF 52.9 163.2 (2.9) 21.8 6.3 (<1) 2.7 0.0 (0.0) 12.0 8.9 (<1) 10.6 3.8 (<1) BT 3.8 63.7 (8.3) 58.3 86.6 (<1) 19.3 6.2 (<1) 18.4 18.5 (<1) <1.0 0.3 (<1) EL 90.3 162.6 (2.5) 9.7 15.7 (2.3) 0.0 0.0 (0.0) 0.0 0.0 (0.0) 0.0 0.0 (0.0) HD 92.7 3.1 (<1) 5.7 139.3 (83.2) 1.7 40.2 (82.0) 0.0 0.0 (0.0) 0.0 0.0 (0.0)

b. 0-25% 25-50% 50-75% 75-100% >100% Sit % of Area (m2) (% of % e site Patch Occupied) Area Area (%) % Area Area (%) % Area Area (%) % Area Area (%) DR 47.9 4.0 (<1) 29.9 0.0 (0.0) 8.9 1.1 (<1) 11.7 1.4 (<1) 1.5 0.0 (0.0) HF 52.9 25.4 (<1) 21.8 0.1 (<1) 2.7 0.0 (0.0) 12.0 26.6 (2.1) 10.6 1.9 (<1) BT 3.8 0.0 (0.0) 58.3 12.5 (<1) 19.3 52.2 (1.3) 18.4 26.8 (<1) <1.0 1.9 (1.1) EL 90.3 37.4 (<1) 9.7 1.3 (1.8) 0.0 0.0 (0.0) 0.0 0.0 (0.0) 0.0 0.0 (0.0) HD 92.7 255.3 (9.3) 5.7 1.4 (<1) 1.7 0.0 (0.0) 0.0 0.0 (0.0) 0.0 0.0 (0.0)

Summation of Important Points

Sample seasons for this study found the first year to be a dry year, and the second a wet year. Precipitation totals found significantly less water in the less-disturbed sites watershed compared to the disturbed and restored sites watershed. As such, disturbed and restored sites had higher moisture values compared to less-disturbed. However, increased connectivity at the restored site provided higher wetness values compared to the disturbed.

Decreased connectivity was found to relate to higher moisture heterogeneity at the disturbed sites compared to the restored and less-disturbed sites.

The restored site had intermediate total and exotic richness compared to disturbed and less-disturbed sites. This pattern also held across moisture categories. Across sites, species tended to be PG and PF plant type categories, and OBL and FAC wetland categories.

Species richness and average percent total area estimations (total, plant-type, wetland category, status) decreased between sample years for all sites. Although BT and less- disturbed sites had less exotics they had higher exotic cover values. This was particularly apparent in wet years by PHAQ (Phalaris aquatica, Harding grass).

CHAPTER IV

DISCUSSION

Disturbance in Wet Montane Meadows

Current literature indicates strong relationships between hydrologic disturbances and wetland degradation (Alvarez-Cobelas et al 2001, Wright et al 2003, Loheide 2007,

2009, Marini et al. 2007). Thus, restoration to such damaged systems should first focus on hydrologic reconstruction (Martin and Chambers 2002, Seabloom 2003, Klimkowska et al.

2007). Other important aspects to restoration plans are post-restoration surveys that assess the ecological trajectory (habitat health) of the system in question once hydrologic connection has been reestablished (Zelder and Callaway 1999, SER 2005). While there have been a number of studies on restored habitats, there have been few attempts at restoration through hydrologic reconstruction in wet meadows of the Sierra Nevada and even fewer of those with post-restoration assessment, especially several years post restoration efforts.

This research focused on assessing the ecological trajectory of a restored montane meadow seven years following the reestablishment of its hydrologic connectivity. It was hypothesized that the ecological health of the wet meadow would have moved away from the

“degraded state” (Hobbs and Mooney 1993) based on reestablishing the hydrologic connectivity. The assessment was made by comparing the restored montane meadow to two hydrologically disturbed and two less-hydrologically disturbed meadows in two different watersheds, which could potentially introduce greater variation for assessment.

Comparison Issues with Less- Disturbed Sites

Ideally, the comparison of the restored meadow to the less-disturbed meadows would help indicate how closely the restored site’s moisture values and hydrologic continuity line-up (SER 2005). Substantial efforts were made to identify comparable less-disturbed sites in the same watershed as the restored site, however, no less-disturbed meadows with all necessary attributes could be identified. Therefore, I tried to identify sites in relatively close proximity to the Calaveras watershed that were comparable. However, even with this effort, the less-disturbed sites surveyed in the Upper Merced watershed received less precipitation and were not comparable hydrologically to those in the Upper Calaveras watershed. The less- disturbed sites are fed indirectly via the Merced River while the restored and disturbed sites are fed directly via seasonal surface water. Water at the less-disturbed sites therefore do not flow through them but rather rises through groundwater as a seasonal result of increasing flow in the Merced River (Loheide et al 2009). Thus, as the Upper Merced’s watershed hydrologic regime was so different, and considering moisture’s influence on vegetation patterns (Wright et al 2003, Marini et al 2007, Loheide et al 2009), it was difficult to make meaningful comparisons between meadows in these two watersheds. As such, the assessment for ecological trajectory for the restored meadow will focus more heavily on comparisons with disturbed sites.

Summary of Key Findings

The restored site had the highest moisture values and highest density of wet patches compared to both disturbed and less-disturbed sites. When compared to the disturbed

57 sites, it had less extreme moisture differences between adjacent patches, typical for healthy

58 montane meadows (Ratlif 1985, Ramstead et al 2012, Martin 2002). Also, general patterning of dominant vegetation types within patches for the restored site was much more in line with montane meadow definitions (Fites-Kaufman et al 2007, Sawyer, Wolf, and Evans 2009), especially across the moisture categories, compared to the disturbed sites. As such, the overall hydrologic patterns and vegetation at the restored site indicate a positive ecological trajectory (restoration success) compared to the disturbed meadows.

The restored sites relatively high moisture and connectivity lead to overall high richness values, especially of species one would expect to see in montane meadows relative to plant types and wetland affinities (Ratlif 1985, Murray et al 2003, Boulton 2005). Across sample years all sites saw fluctuations in richness and diversity of wetland species, however the restored site generally was found to have greater resilience most likely owing to its increased moisture and connectivity. Regarding exotic species, the restored site had higher relative areas than the other sites, though they were still represented a small relative total area for the site. Again, abiotic characteristics distinguished the restored site and continued to drive biotic patterns in ways distinctive of the disturbed and less-disturbed sites indicating restoration success, though areas of exotics was concerning.

Restoration Assessment

As there is no cookbook for restoration planning, each unique wetland system needs its own unique restoration methods, yet restoring hydrology or rewetting of the soil is considered the most important aspect of any wetland restoration project (Martin and

Chambers 2002, Bodegom 2006, Klimkowska et al. 2007). However, only focusing on

abiotic factors is not enough, as might be the case for BT where an attempt to restore the 58

59 hydrology was the extent of the restoration effort, there must also be a biotic component focusing on native plant propagation and invasive species control (Klimkowska et al. 2007).

Donath (2003) found that restoration success is influenced by proximity of native remnant stands. There were some remnant stands of intact montane meadow species at the restored site in the original assessment before the restoration (Boyd and Woodward 1988), and it was those parts of the meadow that appear to have made the most positive ecological trajectory.

The fact that even parts of the disturbed meadows in this study have portions representative of intact montane meadow systems indicates that restoration to them has a higher potential for success. It is important to note that characteristic wetland plants do not always dominate newer restorations right away as it takes time for hydrology to adjust after restoration (Meyer et al. 2010). This being the case, models could be used to design and predict vegetation patterning based on potential changes to the hydrology (Loheide and Gorelick 2007). This would be especially helpful since this restoration is still new and needs continual monitoring/management of the hydrology until it is more sustainable.

Follow up assessments are also imperative, and choice of reference sites strongly affects the outcome of a comparison to evaluate restoration success (Kentula 2000, Morgan and Short 2002). As discovered in this study, although intensive research was done to find appropriate reference sites, the chosen less-disturbed sites still ended up being inappropriate for comparison. Comparison of the restored site to historical (pre-industrial revolution) ecosystems is hardly achievable given increased human population with accompanying levels of pollution, landscape fragmentation, and species extinctions (Wolters et al. 2005). Still, there must be some way of assessing ecological trajectories for the restored site beyond

59 comparing them to disturbed sites as in this study. This study emphasizes the need to

60 understand the hydrologic connectivity and ensure that sites that establish the basis for comparisons are appropriate. There has been some research in using species-by-habitat data matrices, generated by biodiversity surveys, to evaluate outcomes of restoration when no reference sites exist (Brewer and Menzel 2009). In studies like this one, a solid next step might be looking into a model-based methodology as described above that can help further distinguish the ecological trajectory of restorations similar to the restored site in this study. It also might be worth using more sensitive measurements to better accurately quantify the trajectory of BT. Some examples are as follows:

 Redox potential: This measurement is important for determining vegetation patterns

(Purdy and Moyle 2006), and linked with species richness and total plant cover (Dwire et al.

2006).

 Channel incision rate, direction (positive/negative), and stability: This type of measurement is especially important for the small channel still present in BT as well as future reference if disturbed sites are ever restored (Micheli and Kirchner 2002).

 Soil carbon: Relates to function within wet meadows, this is especially important when comparing hydrologically disturbed meadows (Norton et al. 2011, Pope et al. 2015).

Another item crucial for successful restoration and restoration upkeep is supportive management planning (Seabloom 2003). One of the most important things, at both

BT and other restoration sites, is dealing with any potential local anthropogenic events that could turn/slow the trajectory. First, I would suggest continual monitoring of the hydrology focusing on connectivity and channelization. I would also suggest taking steps to potentially prevent any hydrological damage, especially in relationship to trails (Scott 1998). At BT this

60 means firmly designating trail usage onto the boardwalks, re-routing any trails that lead to

61 hydrologically sensitive areas, and restoration/reclamation of previously used trails by visitors (Eagan et al 2000, 2004). Establishing a photo-monitoring program could be helpful for monitoring potential channeling due to trails as well as conifer encroachment (Vale

1987). Monitoring of exotic species is particularly important at the restored site due to the observed areas of exotics. As instances relating to its exotic species spread correlate to heavy foot traffic (Eagan et al 2000, 2004), trail work becomes even more important. Potentially,

BT could set up boot cleaning stations by major trailheads and/or create an annual volunteer/citizen study projects to help rid exotics from the meadow and/or make a display in the visitor center discussing what visitors can do to help deter exotic establishment and spread.

Existing literature indicates that general post-project monitoring tends to be short- term, so while the project may show a positive ecological trajectory, how the restored landscape functions within the natural variability of the system over long term is not well understood (Pope et al. 2015). That being said, there has been success with other wetland meadow restorations (Martin and Chambers 2002, Billeter et al. 2007, Poptcheva et al. 2009) but it is important to note that restorations rarely follow linear recovery. There have been some restorations and research more locally to the Sierra Nevada (Egan et al. 2000, 2004), however they are few and far between. That leaves many Sierra Nevada meadows potentially in need of assessment, monitoring, and potential restoration.

So what is the ecological trajectory of the restored meadow at Big Trees State

Park? Positive factors: High soil moisture levels and hydrologic connectivity, richness and diversity values definitive of wet montane meadows, and little conifer encroachment. Factors

61 of concern: Exotic areas throughout the meadow, especially across dry soil moisture zones,

62 and the presence of a small channel. Overall, the restored site has many more attributes indicating a positive ecological trajectory signifying that the rewetting techniques applied were successful. However, it is clear that adaptive management is needed to further monitor this meadow to adjust the restoration plan as necessary depending on if any of the concerning observed trends begin to slow/decrease this sites overall positive trajectory. To conclude,

John Muir described Sierra Nevada meadows as being “so complete you cannot see the ground.” However, if California continues to have drought conditions into an unpredictable future, there may be negative ecological trajectories for all Sierra Nevada montane meadow systems despite their hydrological status.

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APPENDIX A

Plant list. Accumulated from 2011 and 2012 field seasons Water Plant Species Symbol Family Status affinity type Camassia quamash CAQU2 Agavaceae N FACW PF Allium sanbornii ALSA Alliaceae N NI PF Angelica breweri ANBR5 Apiaceae N NI PF Perideridia lemmonii PELE5 Apiaceae N NI PF Cicuta douglasii CIDO Apiaceae N OBL PF Heracleum maximum HEMA80 Apiaceae N FACU PF Asarum lemmonii ALSE Aristolochiaceae N OBL PF Solidago canadensis SOCA6 Asteraceae N FACU PF Senecio integerrimus SEIN2 Asteraceae N FAC PF Achillea millefolium ACMI2 Asteraceae N FACU PF Senecio clarkianus SECL Asteraceae N FACW PF Helenium bigelovii HEBI Asteraceae N OBL PF Anaphalia margaritacea ANMA Asteraceae N NI PF Artemisia ludoviciana ssp incompta ARLUI2 Asteraceae N FACU PF Madia gracilis MAGR3 Asteraceae N NI AF Symphyotrichum bracteolatum SYBR2 Asteraceae N FAC PF Jensia rammii JERA Asteraceae N NI AF Erigeron inornatus var. inornatus ERINI Asteraceae N UPL PF Hypochaeris radicata HYRA3 Asteraceae I FACU PF Cirsium vulgare CIVU Asteraceae I FAC PF Tragopogon dubius TRDU Asteraceae I NI PF Alnus rhombifolia ALRH2 Betulaceae N FACW T/S Nemophila maculata NEMA Boraginaceae N NI AF Myosotis discolor MYDI Boraginaceae NN NI AF Capsella bursa-pastoris CABU2 Brassicaceae NN FAC AF

Stellaria graminea STGR Caryophyllaceae NN UPL PF

Plant list cont. Water Plant Species Symbol Family Status affinity type Chenopodium album CHAL7 Chenopodiaceae NN FAC PF Carex nebrascensis CANE2 Cyperaceae N OBL PG Carex nuda CANU5 Cyperaceae N FACW PG Carex feta CAFE4 Cyperaceae N OBL PG Scirpus microcarpus SCMI2 Cyperaceae N OBL PG Carex pachystachya CAPA14 Cyperaceae N FACW PG Carex vesicaria CAVE6 Cyperaceae N OBL PF Carex aurea CAAU3 Cyperaceae N OBL PG Carex serratodens CASE2 Cyperaceae N FACW PG Carex utriculata CAUT Cyperaceae N NI PG Carex lenticularis CALE8 Cyperaceae N OBL PG Carex praegracilis CAPR5 Cyperaceae N FACW PG Eleocharis parishii ELPA4 Cyperaceae N FACW PG Carex gracilior CAGR9 Cyperaceae N NI PG Carex athrostachya CAAT3 Cyperaceae N FACW PG Carex echinata CAEC Cyperaceae N OBL PG Equisetum hyemale EQHY Equisetaceae N FACW PF Equisetum arvense EQAR Equisetaceae N FAC PF Vaccinium cespitosum VACE Ericaceae N NI T/S Rhododendron occidentale RHOC Ericaceae N FAC T/S Hosackia oblongifolia var. oblongifolia LOOBO2 Fabaceae N OBL PF Lupinus latifolius var. columbianus LUAL4 Fabaceae N FAC PF Trifolium hirtum TRHI4 Fabaceae N NI AF Acmispon americanus var. americanus ACAM Fabaceae N UPL AF

Trifolium dubium TRDU2 Fabaceae NN FACU AF 75

Vicia sativa ssp. nigra VISAN2 Fabaceae NN FACU AF

Plant list cont. Water Plant Species Symbol Family Status affinity type Lathyrus latifolius LALA4 Fabaceae NN OBL PF Centaurium erythraea CEER5 Gentianaceae I FAC AF Hypericum anagalloides HYAN2 Hypericaceae N OBL A/PF Hypericum formosum HYFO7 Hypericaceae N NI PF Iris hartwegii IRHA Iridaceae N NI PF Juncus macrandrus JUMA2 Juncaceae N OBL PG Juncus ensifolius JUEN Juncaceae N FACW PG Juncus effusus JUEF Juncaceae N FACW PG Juncus nevadensis JUNE Juncaceae N FACW PG Mentha arvensis MEAR4 Lamiaceae N FACW PF Prunella vulgaris PRVU Lamiaceae N FAC PF Agastache urticifolia AGUR Lamiaceae N FACW PF Pycnanthemum californicum PYCA Lamiaceae N NI PF Mentha spicata MESP3 Lamiaceae NN OBL PF Lilium parvum LIPA3 Liliaceae N OBL PF Limnanthes alba ssp. alba LIALA Limnanthaceae N FACW AF Limnanthes alba LIAL3 Limnanthaceae N FACW AF Sidalcea calycosa ssp. Calycosa SICAC3 Malvaceae N OBL AF Veratrum californicum var. californicum VECAC2 Melanthiaceae N OBL PF Mollugo verticilla MOVE Molluginaceae NN FAC AF Null Null Null Null Null T/S Null Null Null Null Null F Nuphlar polysepala NULUP Nymphaeaceae N OBL PF Epilobium ciliatum EPCI Onagraceae N FACW PF Epilobium foliosum EPFO Onagraceae N NI PF

Clarkia amoena CLAM Onagraceae N NI PF

Plant list cont. Water Plant Species Symbol Family Status affinity type Platanthera dilatata var. leucostachys PLDIL Orchidaceae N FACW PF Spiranthes romanzoffiana SPRO Orchidaceae N OBL PF Mimulus primuloides MIPR Phrymaceae N OBL PF Mimulus laciniatus MILA2 Phrymaceae N FAC AF Mimulus floribundus MIFL2 Phrymaceae N OBL AF Mimulus guttatus MIGU Phrymaceae N FACW A/PF Mimulus torreyi MITO Phrymaceae N NI AF Mimulus bicolor MIBI4 Phrymaceae N FACW AF Pinus ponderosa PIPO Pinaceae N UPL T/S Gratiola ebracteata GREB Plantaginaceae N OBL AF Penstemon rydbergii PERY Plantaginaceae N FAC PF Veronica americana VEAM2 Plantaginaceae N OBL PF Plantago lanceolata PLLA Plantaginaceae I FAC PF Eragrostis pectinacea var. pectinacea ERPEP2 Poaceae N FAC AG Poa howellii POHO6 Poaceae N NI AG Vulpia microstachys VUMI Poaceae N NI AG Festuca microstachys FEMI2 Poaceae N NI PG Panicum acuminatum PAAC5 Poaceae N FACW PG Elymus glaucus ssp. glaucus ELGLG Poaceae N FAC PG Danthonia californica DACA3 Poaceae N FACW PG Bromus arenarius BRAR3 Poaceae NN NI AG Poa compressa POCO Poaceae NN FAC PG Phalaris aquatica PHAQ Poaceae I FAC PG Anthoxanthum odoratum ANOD Poaceae I FACU PG Dactylis glomerata DAGL Poaceae I FACU PG

Holcus lanatus HOLA Poaceae I FAC PG

Plant list cont. Water Plant Species Symbol Family Status affinity type Bromus hordeaceus BRHO2 Poaceae I FACU AG Collomia heterophylla COHE2 Polemoniaceae N FAC AF Collomia linearis COLI2 Polemoniaceae N FACU AF Rumex acetosella RUCA3 Polygonaceae I FAC PF Rumex crispus RUCR Polygonaceae I FACW PF Dodecatheon hendersonii DOHE Primulaceae N FACU PF Delphinium hansenii DEHA Ranunculaceae N NI PF Ranunculus californicus RACA2 Ranunculaceae N FAC PF Aconitum columbianum ACCO4 Ranunculaceae N FACW PF Aquilegia formosa AQFO Ranunculaceae N FAC PF Rubus ursinus RUUR Rosaceae N FAC T/S Potentilla gracilis POGR9 Rosaceae N FACW PF Drymocallis glandulosa var. glandulosa DRGL7 Rosaceae N FAC PF Fragaria vesca FRVE Rosaceae N UPL PF Malus domestica MAPU Rosaceae NN NI T/S Salix melanopsis SAME2 Salicaceae N FACW T/S Verbascum thapsus VETH Scrophulariaceae I NI PF Triteleia hyacinthina TRHY3 Themidaceae N FACW PF Brodiaea coronaria BRCO3 Themidaceae N FAC PF Typha domingensis TYDO Typhaceae N OBL PF Viola macloskeyi VIMA2 Violaceae N OBL PF

APPENDIX B

USFWS Indicator categories Abbreviation Name Description Obligate OBL Occurs almost always (estimated probability 99%) under natural conditions in wetlands. Wetland Facultative Usually occurs in wetlands (estimated probability 67%-99%), but occasionally found in non- FACW Wetland wetlands. FAC Facultative Equally likely to occur in wetlands or non-wetlands (estimated probability 34%-66%). FACU Facultative Usually occurs in non-wetlands (estimated probability 67%-99%), but occasionally found on Upland wetlands (estimated probability 1%-33%).

UPL Occurs in wetlands in another region, but occurs almost always (estimated probability 99%) under Obligate natural conditions in non-wetlands in California. If a species does not occur in wetlands in any Upland region, it is not on the National List.

NA No agreement The regional panel was not able to reach a unanimous decision on this species. NI No indicator Insufficient information was available to determine an indicator status

APPENDIX C

Patch classification and characteristics across sites for 2012 field season Sp.- Wetness Size Avg. Percent Site Patch Name Type Category (m^2) Richness NV DR SOCA6 PF 0-25 122.1 9 55.9 DR CAQU2 PF 25-50 233.2 20 28.6 DR VECAC2 #1 PF 75-100 144.8 9 29.9 DR VECAC2 #2 PF 75-100 140.2 11 11.7 DR VECAC2 #3 PF 0-25 229.2 13 36.5 DR CANE2 #1 PG 25-50 607.2 10 10.0 DR CANE2 #2 PG 50-75 48.3 7 21.8 DR CANE2 #3 PG 50-75 152.3 7 14.2 DR CANE2 #4 PG 25-50 371.7 9 18.9 DR CANE2 #5 PG 0-25 327.6 16 18.1 DR JUMA2 #1 PG 25-50 110.9 11 25.3 DR JUMA2 #2 PG 50-75 179.0 17 45.4 DR JUMA2 #3 PG 50-75 122.4 18 62.1 DR MS JUMA2/JUEF MS 50-75 61.4 12 37.6 DR ALSA #1 PF 75-100 557.7 31 53.7 DR ALSA #2 PF 75-100 29.3 11 44.0 DR SECL #1 PF >100 115.2 17 35.3 DR SECL #2 PF 50-75 15.7 4 59.0 DR SECL #3 PF 50-75 30.3 6 23.5 DR JUEF #1 PG 50-75 53.2 11 42.0 DR JUEF #2 PG 25-50 121.2 19 43.1 DR SICAC3 AF 0-25 200.5 25 38.2 DR MS SOCA6 MS 25-50 772.0 32 52.5 DR MS #1 MS 0-25 288.3 37 35.1 DR MS #2 MS 0-25 349.5 28 56.2 82

DR MS CAGR9/ERPEP2 #1 MS 0-25 248.1 12 36.8

Patch classification and characteristics across sites for 2012 field season cont. Wetness Size Avg. Percent Site Patch Name Sp.-Type Category (m^2) Richness NV MS CAGR9/ERPEP2 DR #2 MS 0-25 1061.3 14 28.4 MS CAGR9/ERPEP2 DR #3 MS 0-25 723.4 24 37.2

HF VECAC2 PF 50-75 34.9 5 1.0 HF SCMI2 PG 0-25 46.8 7 8.3 HF SICAC3 AF 0-25 61.8 7 41.0 HF SYBR2 PF 0-25 173.6 14 33.8 HF TYDO PF >100 940.2 7 47.0 HF SOCA6 #1 PF 0-25 41.9 24 44.4 HF SOCA6 #2 PF 0-25 110.7 9 45.0 HF SOCA6 #3 PF 25-50 125.3 15 42.0 HF SOCA6 #4 PF 0-25 107.4 17 29.2 HF SOCA6 #5 PF 0-25 521.2 17 24.1 HF SOCA6 #6 PF 0-25 144.7 14 29.8 HF SOCA6 #7 PF 0-25 307.2 8 29.0 HF LUAL4 #1 PF 25-50 438.0 6 7.1 HF LUAL4 #2 PF 0-25 80.7 7 31.0 HF LUAL4 #3 PF 25-50 251.2 6 16.8 HF MS LUAL4/SICAC3 MS 0-25 437.7 7 2.1 HF JUMA2 #1 PG 50-75 44.9 13 26.8 HF JUMA2 #2 PG 25-50 332.1 23 16.6 HF JUEF #1 PG 25-50 20.1 8 33.8 HF JUEF #2 PG 25-50 68.8 13 28.8 HF JUEF #3 PG 50-75 127.7 5 35.3

HF JUEF #4 PG 75-100 189.1 5 14.5 83

HF JUEF #5 PG 25-50 1079.4 15 17.9

Patch classification and characteristics across sites for 2012 field season cont. Wetness Size Avg. Percent Site Patch Name Sp.-Type Category (m^2) Richness NV HF CAUT #1 PG 75-100 279.1 27 24.4 HF CAUT #2 PG 75-100 335.1 20 21.4 HF CAUT #3 PG 50-75 27.2 9 13.8 HF CALE8 #1 PG 75-100 42.2 4 23.0 HF CALE8 #2 PG 50-75 49.1 6 5.5 HF RACA2 #1 PF 75-100 265.9 22 48.6 HF RACA2 #2 PF 75-100 28.4 14 22.0 HF RACA2 #3 PF 75-100 136.5 9 6.8 HF CAFE4 PG >100 192.8 8 10.0 HF MS RUCA3 MS 0-25 500.4 24 66.4 HF MS MS 0-25 2634.9 41 42.9 HF MS CAGR9 #1 MS 0-25 406.8 27 46.8 HF MS CAGR9 #2 MS 0-25 51.9 9 27.5 BT CANE2 PG 75-100 522.9 10 37.7 BT SCMI2 PG 75-100 136.3 10 21.0 BT JUEF PG 75-100 301.8 15 56.8 BT TRHI4 AF 50-75 438.8 16 38.2 BT TYDO PG >100 71.5 8 60.5 BT VECAC2 #1 PF 75-100 711.9 6 22.3 BT VECAC2 #2 PF 75-100 495.3 23 32.5 BT VECAC2 #3 PF 75-100 182.0 1 12.0 BT VECAC2 #4 PF 75-100 52.5 4 17.5 BT SOCA6 #1 PF 50-75 392.9 19 30.4 BT SOCAG #10 PF 0-25 122.4 11 57.4 BT SOCA6 #2 PF 25-50 238.5 15 47.6

BT SOCA6 #3 PF 25-50 143.9 14 51.0

Patch classification and characteristics across sites for 2012 field season cont. Wetness Size Avg. Percent Site Patch Name Sp.-Type Category (m^2) Richness NV BT SOCA6 #4 PF 25-50 53.5 13 50.5 BT SOCA6 #5 PF 25-50 154.6 15 50.6 BT SOCA6 #6 PF 25-50 144.4 19 45.2 BT SOCA6 #7 PF 25-50 80.5 10 47.5 BT SOCA6 #8 PF 25-50 52.7 6 34.0 BT SOCA6 #9 PF 25-50 174.0 6 52.3 BT POGR9 #1 PF 50-75 1270.2 24 28.3 BT POGR9 #2 PF 25-50 190.9 18 33.0 BT LUAL4 #1 PF 50-75 142.8 22 35.5 BT LUAL4 #2 PF 50-75 174.8 14 36.3 BT LUAL4 #3 PF 25-50 981.8 17 25.5 BT LUAL4 #4 PF 25-50 216.4 6 36.5 BT LUAL4 #5 PF 25-50 304.2 7 22.8 BT JUMA2 #1 PG 50-75 114.2 5 42.0 BT JUMA2 #2 PG >100 104.3 20 28.1 BT JUMA2 #3 PG 75-100 132.9 14 40.2 BT JUMA2 #4 PG 75-100 47.1 9 30.0 BT JUMA2 #5 PG 75-100 97.8 16 27.9 BT CAFE4 #1 PG 75-100 317.8 14 36.7 BT CAFE4 #2 PG 50-75 566.3 6 42.7 BT ANOD #1 PG 0-25 88.3 12 59.4 BT ANOD #2 PG 0-25 127.1 16 53.3 BT ANOD #3 PG 0-25 426.5 13 54.0 BT MS #1 MS 75-100 325.6 19 54.3 BT MS #2 MS 75-100 58.5 9 43.5 85

BT MS #3 MS 75-100 164.6 16 38.8

Patch classification and characteristics across sites for 2012 field season cont. Wetness Size Avg. Percent Site Patch Name Sp.-Type Category (m^2) Richness NV BT CAGR9 #1 PG 75-100 169.5 4 48.7 BT CAGR9 #2 PG 25-50 5832.4 30 41.8 BT CAGR9 #3 PG 25-50 3093.3 21 52.7 BT CAGR9 #4 PG 50-75 753.3 14 48.6 EL LOOBO2 PF 25-50 602.3 20 42.4 EL RUCA3 PF 0-25 113.7 17 61.7 EL CAUT PG 0-25 96.8 5 61.3 EL MEAR4 PF 0-25 599.9 12 51.6 EL CANE2 #1 PG 0-25 1004.5 15 42.3 EL CANE2 #2 PG 0-25 56.1 8 37.3 EL CANE2 #3 PG 25-50 88.7 9 56.4 EL POHO6 #1 AG 0-25 419.9 12 63.9 EL POHO6 #2 AG 0-25 178.8 12 55.4 EL MS MEAR4/POHO6 MS 0-25 407.8 23 54.3 EL MS POHO6 MS 0-25 1128.9 23 56.1 EL MS #1 MS 0-25 1454.0 20 67.0 EL MS #2 MS 0-25 969.1 27 65.7 HD MEAR4 PF 0-25 71.1 10 44.2 HD ARLUI2 PF 0-25 90.5 9 48.8 HD ELGLG PG 0-25 95.1 9 71.5 HD ERPEP2 #1 AG 0-25 138.2 4 73.0 HD ERPEP2 #2 AG 0-25 90.7 8 38.0 HD SCMI2 #1 PG 0-25 86.9 3 21.7 HD SCMI2 #2 PG 0-25 218.6 6 20.5 HD SCMI2 #3 PG 0-25 89.7 4 29.0

HD SCMI2 #4 PG 0-25 113.9 13 53.5

Patch classification and characteristics across sites for 2012 field season cont. Wetness Size Avg. Percent Site Patch Name Sp.-Type Category (m^2) Richness NV HD HOLA #1 PG 50-75 7.1 2 15.2 HD HOLA #2 PG 50-75 7.5 0 0.2 HD HOLA #3 PG 50-75 34.4 2 17.0 HD HOLA #4 PG 25-50 48.5 2 0.0 HD HOLA #5 PG 25-50 119.0 5 19.8 HD POCO #1 PG 0-25 137.1 12 50.6 HD MS MS 0-25 811.7 14 37.2 HD MS POCO #1 MS 0-25 250.6 5 32.3 HD MS POCO #2 MS 0-25 541.0 12 42.8

APPENDIX D

Number of patches at each site dominated by different plant types (AF=annual forb, AG=annual graminoid, PF=perennial forb,PG=perennial graminoid, MS=multiple species) along with the total area (m2) and (relative area) of the site the different patch types encompass. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. AG AF PG PF MS Area Patch Area Patch Area Patch Area Patch Area Patch Site (%Area) Count (%Area) Count (%Area) Count (%Area) Count (%Area) Count DR 0.0 (0.0) 0 200.5 (2.7) 1 2093.9 (28.2) 10 1617.6 (21.1) 10 3504.1 (47.2) 7 HF 0.0 (0.0) 0 61.8 (<1) 1 2834.2 (26.6) 14 3708.1 (34.9) 16 4031.7 (37.9) 5 BT 0.0 (0.0) 0 438.8 (2.2) 1 12903.3 (64.5) 18 6279.9 (31.1) 21 548.8 (2.7) 3 EL 598.7 (8.4) 2 0.0 (0.0) 0 1246.1 (17.5) 4 1315.9 (18.5) 3 3959.8 (55.6) 4 HD 228.9 (7.8) 2 0.0 (0.0) 0 957.9 (32.4) 11 161.7 (5.5) 2 1603.3 (54.3) 3

APPENDIX E

Tables a-b. Areas (m2) of plant type (a) and plant wetland affinities (b) by soil moisture categories for each site in 2012 field season. a. Plant type and soil moisture category species areas by site Plant Type AG AF AF/PF PG PF T/S Site Moisture category 0-25% 112.7 407.8 3.9 938.6 290.0 10.4

25-50% 25.3 99.7 0.0 775.6 306.5 0.0

DR 50-75% 0.0 28.5 8.7 121.3 27.1 0.0

75-100% 6.0 61.1 4.2 123.1 222.8 0.0

>100% 1.2 10.4 0.0 9.2 48.4 0.0

0-25% 124.2 608.8 0.0 787.9 1085.1 0.0

25-50% 7.1 263.5 0.0 421.9 976.1 0.0

HF 50-75% 0.3 2.4 0.0 48.9 38.3 0.0

75-100% 0.9 94.0 0.0 407.6 196.6 0.0

>100% 3.9 64.0 0.0 229.2 367.0 0.0

0-25% 21.2 3.3 0.0 103.2 297.2 0.0

25-50% 962.5 93.5 0.0 1440.5 2751.3 0.0

BT 50-75% 128.0 172.7 0.0 439.5 1318.9 0.0

75-100% 76.1 23.5 0.0 608.7 948.4 0.0

>100% 0.6 2.1 7.1 66.3 21.4 0.0

0-25% 572.5 13.3 0.6 859.3 474.5 0.6 EL 25-50% 52.9 0.7 0.0 71.0 165.9 0.0

0-25% 597.4 1.7 0.0 744.9 327.5 2.1

HD 25-50% 0.0 0.0 0.0 142.8 1.2 0.0

50-75% 0.0 0.0 0.0 41.5 0.0 0.0

b. Wetland classification and soil moisture category species areas by site Wetland Classifications FAC OBL FAC FACU UPL NI Site Moisture category W 0-25% 1231.6 28.8 90.8 138.7 3.4 270.1

25-50% 760.5 165.3 52.0 94.2 1.5 133.6

DR 50-75% 144.4 25.2 2.1 7.5 0.0 6.3

75-100% 238.0 54.3 10.9 25.1 0.0 88.8

>100% 12.7 34.6 3.5 11.5 0.0 6.9

0-25% 704.9 262.2 616.4 502.3 13.3 506.9

25-50% 355.6 312.7 860.3 82.8 0.0 57.0

HF 50-75% 76.5 0.0 4.8 0.3 0.0 8.0

75-100% 189.9 132.5 196.8 4.7 0.0 175.2

>100% 435.7 61.1 153.8 1.9 0.0 11.6

0-25% 11.4 37.5 159.6 186.9 0.0 29.3

25-50% 247.3 535.6 1196.6 1325.5 20.6 1909.7

BT 50-75% 675.3 364.0 205.5 320.0 25.8 468.6

75-100% 790.4 81.6 210.0 316.3 13.6 244.9

>100% 64.9 19.2 3.0 8.8 1.0 0.6

0-25% 448.9 259.8 366.3 61.5 4.5 781.0 EL 25-50% 179.3 33.1 20.0 1.2 0.0 56.8

0-25% 418.6 141.5 895.9 175.4 1.4 40.9

HD 25-50% 0.9 1.2 141.9 0.0 0.0 0.0

50-75% 0.7 0.0 33.7 0.0 0.0 0.0

APPENDIX F

Percentage of site area in each soil moisture category and total area of non-vegetated (NV) ground (m2) and the (percent area that comprises each moisture category) for 2012. Sites are listed along a disturbance gradient with DR and HF categorized as disturbed, BT as restored, and EL and HD categorized as least disturbed. For example, in DR 47.9% of total site is in the 0-25% moisture category, and there was a total of 1786.5m2 of non-vegetated area that represents 50.3% of the total area within the 0-25% moisture category. 0-25% 25-50% 50-75% 75-100% >100% NV Area NV Area NV Area NV Area NV Area Site % Area (%) % Area (%) % Area (%) % Area (%) % Area (%) DR 47.9 1786.5 (50.3) 29.9 895.9 (40.4) 8.9 217.4 (32.8) 11.7 454.7 (52.1) 1.5 46.1 (40.0)

HF 52.9 3020.7 (53.7) 21.8 663.2 (28.6) 2.7 13.2 (4.6) 12.0 414.3 (32.5) 10.6 468.9 (41.4)

BT 3.8 339.5 (44.4) 58.3 6239.0 (53.5) 19.3 1235.2 (32.1) 18.4 1753.5 (47.2) <1.0 78.2 (44.5)

EL 90.3 4500.4 (70.0) 9.7 400.5 (57.9) 0.0 0.0 (0.0) 0.0 0.0 (0.0) 0.0 0.0 (0.0)

HD 92.7 1113.1 (40.7) 5.7 23.5 (14.0) 1.7 6.9 (14.1) 0.0 0.0 (0.0) 0.0 0.0 (0.0)