FLORISTIC AFFINITIES OF THE SAN JOAQUIN ROADLESS AREA,

IΝΥΟ NATIONAL FOREST, MONO COUNTY, CALIFORNIA

by

Helen M. Constantine-Shull

A Thesis

Presented to

The Faculty of Humboldt State University

In Partial Fulfillment

of the Requirements for the Degree

Master of Arts

In Biology

May, 2000 FLORISTIC AFFINITIES OF THE SAN JOAQUIN ROADLESS AREA,

INΥΟ NATIONAL FOREST, MONO COUNTY, CALIFORNIA

by

Helen M. Constantine-Shul1

Approved by the Master's Thesis Committee:

John O. Sawyer Jr., Major Professor

Mike Mesler, Committee Member

Ken Lang, Committee Member

Robert Curry, Committee Member

Milton Boyd, Graduate Coordinator

Ronald A. Fritzsche, Dean for Research and Graduate Studies ABSTRACT

I surveyed the flora of the San Joaquin Roadless Area, a 44km2 area on the east side of the central Sierra, and then analyzed its floristic affinities (origins) using Sorensen's similarity coefficient, cluster analysis, and Bray-Curtis ordination. The analyses compare the area's flora to 12 other California mountain floras, and suggest an origin for recolonizing taxa after severe disturbances by Holocene volcanic activity. I have described the historic and present climate, geology, physiography, and vegetation of the area for a better understanding of the origin and condition of the flora.

The San Joaquin Roadless Area flora consists of 446 taxa, including two rare

(CLAPS List 1Β) plants, Lupinus duranii Eastw. and Arabis pinzlae Rollins. The ranges of

3 taxa were extended. The flora is centered in geographic location, species composition, and proportion of phytogeographic elements in relation to other California mountain floras.

Its species composition is most similar to Tuolumne Meadows (56%) and vicinity and least similar to the San Bernardino Mountains (30%). The average similarity between all 13 floras examined is 39%.

The cluster analysis suggests three major groups of floras, 1) Sierran and Westside floras, 2) Great Basin and eastside floras, and 3) floras not similar to either group. My flora is included with other Sierran and Westside floras. The Bray-Curtis ordination shows a similar pattern, but with the three axes accounting for only 29% of the variation.

Ordinations on subsets of the floras (1: Floras of similar size, 2: Floras close to the San

Joaquin Roadless Area, and 3: Sierran floras) also show a similar pattern, with the three axes accounting for >70% of the variation between the floras.

The phytogeographic affinities of the entire San Joaquin Roadless Area are generally consistent with those of the entire Sierran flora. However, within each affinity category they differ. The San Joaquin Roadless Area contains a higher percentage of Old

iii Cordilleran taxa, and lower percentages of Circumboreal, Northern Hemisphere, Lowland

California, and Great Basin taxa. A cluster analysis of the percentages in each affinity category shows a similar pattern to that of the other analyses.

Conflicting extremes of wet and dry environments, due to weather patterns and volcanic soils, have come together in the San Joaquin Roadless Area to create a moderate environment that supports a newly immigrated, centrally aligned flora. The availability of favorable conditions has influenced the relative success of colonizing taxa with different origins and adaptations. The proportions of taxa from each region of origin are reflected in the dominating moisture regimes of the area.

My results confirm the Mammoth Gap as an effective migration corridor for

Lowland California taxa to the east side of the Sierra. Despite destructive volcanic disturbances, there is a possibility of alpine refugia where some of the first colonizers of the study area survived. A Sierran origin for the rare plant, Arabis pinzlae is suggested.

iv ACKNOWLEDGEMENTS

This thesis would not have been possible without the wonderful and generous support of many people. I would like to thank everyone involved for helping me make this project happen. My major professor, John Sawyer, encouraged me through the years with unending patience and perceptive guidance. I will always cherish John's inspiring classes and our journeys to the mountains and deserts. His extensive knowledge and engaging, enthusiastic discussions helped me to better understand the plants and their setting.

My committee members, Mike Mesler, Ken Lang, and Robert Curry provided valuable comments and feedback on the thesis. I am fortunate to have taken Dr. Mesler's plant taxonomy class and taught his laboratory. These gave me a strong foundation for identifying my specimens. Dr. Curry of UC Santa Cruz—CSU Monterey Bay contributed his first hand knowledge of the area and its geology and traveled to the study area to hike with me.

I am especially grateful to Dr. G. Ledyard Stebbins who shared his knowledge on the origins of Sierran plants and determined the phytogeographic affinity of each taxon on my species list. Bruce Bingham of the U.S. Forest Service provided valuable computer assistance with access to the Paradox database and matrix format conversion.

My thanks and appreciation go out to Connie Millar of the U.S. Forest Service for securing funding for this project through the Sierra Nevada Ecosystem Project. Her guidance and knowledge on the paleobotany of the area provided me with insight as to the origins of the area's flora. The Biology Graduate Student Association also provided financial assistance for thesis binding.

Stew Winchester of Diablo Valley College opened my eyes to the world of plant ecology and encouraged me to continue my studies at the graduate level. Dr. Dennis Walker clearly and thoroughly introduced me to botany and gave me the chance to teach and apply

v my art to the science. Sally Miller of Friends of the Ingo was my inspiration for studying the San Joaquin Roadless Area. She also provided many unpublished local references.

The contributions of many others helped make this project a success. Teresa

Sholars of College of the Redwoods, Ft. Bragg and James Morefield of the Nevada

Natural Heritage Program confirmed my rare plant identifications. Gordon Leppig provided plant presses and helped with identifications in the HSU Herbarium. Botanical professionals Dean Taylor and Mark Bagley offered their guidance and information on plants and literature resources. Greg Reis of the Mono Lake Committee introduced me to the world wide web and its many climate data resources. Richard Perloff and Kathleen

Nelson of the U.S. Forest Service issued me collecting permits. My peers, Margaret

Willits, Ann Francis, Stassia Samuels, and Julie Evans among others shared valuable

knowledge and observations with me.

Special thanks go out to all of my friends on the Eastside and at the Mono Lake

Committee for their encouragement, companionship, and places to stay while I was doing

my field work. Tommy Kashirsky provided a warm, dry base camp with protection from

mosquitoes and a large table on which to sort and press my specimens.

Finally, I would like to thank my family whose loving support made all this

possible. My Mother, my fearless and dedicated field partner, kept me company, learned

botany, and added several new plants to my species list. I am forever thankful to my

husband, Richard, for his patience and moral support throughout this project — in the

midst of starting our marriage and our own business. He helped me in the field when he

could get away, and did everything when I was gone or writing for weeks at a time. He

always reminded me of the significance of my work, and encouraged me when the finished

thesis seemed so far away. His parents, Marian and Blaine Shull, helped me keep this

project a top priority. I am thankful for their encouragement and support throughout the

years. And thanks, always, to the mountains.

vi This thesis is dedicated to

Dr. G. Ledyard Stebbins

1906-2000

for his inspiration and contributions to the sciences of evolutionary botany and phytogeography. TABLE OF CONTENTS

Page

ABSTRACT iii

ACKNOWLEDGEMENTS v

LIST OF TABLES xi

LIST OF FIGURES xiii

INTRODUCTION 1

Purpose 2

The study area 4

Location and boundaries 4

Physiography 4

Elevational zones 7

Geology 11

Recent volcanics 14

Glacial history 16

Soils 17

Climate 18

Regional 18

Local 19

Weather data 23

Human impacts 28

Research and botanical collecting history 30

Vegetation 31

Paleovegetation 31

Current vegetation 33

viii TABLE OF CONTENTS (CONTINUED)

METHODS 44

Field methods 44

Specimen identification methods 46

Analysis methods 49

Phytogeographic affinities of the taxa 49

Comparison of my flora to other floras 52

Phytogeographic affinities of the floras 53

RESULTS 57

Floristic analysis 57

Rare and uncommon taxa 57

Range extensions 59

Alien taxa 60

Other noteworthy taxa 60

Noteworthy occurrences on the limestone 61

Noteworthy absences 62

Statistical analysis 62

Comparison of all floras 62

Comparison of floras of similar size 67

Comparison of floras close to the San Joaquin Roadless Area 70

Comparison of Sierran floras 73

Floristic affinities of the San Joaquin Roadless Area 76

DISCUSSION 83

Floristic affinities of the San Joaquin Roadless Area 83

Species composition analyses 83

Analyses of phytogeographic affinities 85

ix TABLE OF CONTENTS (CONTINUED)

Mammoth Gap as a migration corridor 87

Origins of the San Joaquin Roadless Area flora 87

The origin of Arabis pinzlae 88

CONCLUSIONS 92

LITERATURE CITED 93

APPENDIX A. Annotated species list of the San Joaquin Roadless Area 106

APPENDIX B. Environmental variables 138

APPENDIX C. Similarity matrices 139

x LIST OF TABLES

Page

Table 1. Summary of average annual weather data for stations near the San Joaquin Roadless Area 25

Table 2. Locations of important collecting areas 47

Table 3. Phytogeographic elements as defined by Stebbins (1982, 1997 Pers. comm.) 51

Table 4. California mountain floras used in the statistical analyses 55

Table 5. Subsets of the thirteen floras used in the analyses 56

Table 6. Coefficients of determination for the correlations between ordination distances and distances in the original n-dimensional space from the analysis of all floras 66

Table 7. Pearson correlations of environmental variables with ordination axes from the analysis of all floras 66

Table 8. Coefficients of determination for the correlations between ordination distances and distances in the original n-dimensional space from the analysis of floras of similar size 69

Table 9. Coefficients of determination for the correlations between ordination distances and distances in the original n-dimensional space from the analysis of floras close to the San Joaquin Roadless Area 72

Table 10. Coefficients of determination for the correlations between ordination distances and distances in the original n-dimensional space from the analysis of Sierran floras 75

Table 11. Comparison of the phytogeographic affinities of my flora with those of the entire Sierran fiora (Stebbins 1982) 77

Table 12. Comparison of the phytogeographic affinities of my alpine flora with those of the entire Sierran alpine flora (Stebbins 1982) 77

Table 13. Phytogeographic affinities of the San Joaquin Roadless Area by elevation zone 77

Table 14. Number of taxa in each affinity category for California mountain floras, arranged from north to south 81

Table 15. Taxa that occur on the Pumice Plain on Mount St. Helens (Titus et al. 1998) and also in all three elevation zones of the San Joaquin Roadless Area 91

xi LIST OF TABLES (CONTINUED)

Table 16. Environmental variables used as the secondary overlay matrix in the ordinations 138

Table 17. Sorensen's similarity matrix for the species composition of all floras 139

Table 18. Sorensen's similiarity matrix of percentages of taxa in each affinity category 140

xii LIST OF FIGURES

Page

Figure 1. An outline map of California and the Ingo National Forest showing the location of the study area with major highways and towns 9

Figure 2. Topographic map of the study area showing major landmarks, scale = 1:24,000x 40% (MaptechTM 1997) 10

Figure 3. Geologic map showing the major rock types of the study area, adapted from Bailey (1989) 13

Figure 4. Sierra Nevada precipitation, monthly means from east and west slope stations for 58 years (Carpenter 1991) 22

Figure 5. Precipitation and temperature at Gem Lake (elevation 2744 m) based on a 60 year mean (Carpenter 1991) 26

Figure 6. Snowfall totals for Mammoth Mountain Ski Area 1969-1999 27

Figure 7. Glass Creek Meadow and San Joaquin Ridge from the east, July 10, 1995 42

Figure 8. Glass Creek Meadow and White Wing Mountain from the summit of June Mountain, July 16, 1996 42

Figure 9. San Joaquin Ridge from the southern edge of Deadman Pass looking north, August 18, 1995 43

Figure 10. The east-side red fir forest beneath San Joaquin Ridge, looking south from White Wing Mountain, July 13, 1995 43

Figure 11. Cluster analysis of the complete set of floras 64

Figure 12. Ordination of all floras, axes one and two 65

Figure 13. Ordination of all floras, axes one and three 65

Figure 14. Cluster analysis of floras of similar size 68

Figure 15. Ordination of floras of similar size 68

Figure 16. Cluster analysis of floras close to the San Joaquin Roadless Area 71

Figure 17. Ordination of floras close to the San Joaquin Roadless Area 71

Figure 18. Cluster analysis of Sierran floras 74

Figure 19. Ordination of Sierran floras 74

xiii LIST OF FIGURES (CONTINUED)

Figure 20. Percent of floras in each geographic affinity category 80

Figure 21. Cluster analysis of percentages in each affinity category 82

xiv INTRODUCTION

The San Joaquin Roadless Area is a relatively pristine area of wilderness quality. It is situated between two heavily developed recreation centers (the towns of Mammoth Lakes and June Lake) in the eastern high Sierra. Until recent years the area has received little attention from botanists, land managers, and the general public. However, increasing demands for the areas resources over the last two decades have brought the area into the

limelight. San Joaquin Ridge has been designated by the Natural Heritage Survey as a

nationally significant site that is in some danger (Burke et al. 1982); and White Wing

Mountain has been proposed as a Research Natural Area due to its significance as a

paleobotanical site. More recently, the entire roadless area has been the subject of a case study project for implementing U.S. Forest Service ecosystem management policies (Millar

1996). Although much research has been done in the area through that project, this thesis is

the only comprehensive flora and phytogeographic study of the area.

At present the roadless area, which serves as the headwaters of the Owens River,

has experienced minimal human impacts and is home to a great diversity of flora and fauna.

Studies have shown the area to contain populations of several rare animal species including

goshawk, marten, mountain beaver at the southern limit of its distribution (Ivey 1979), and

Yosemite toad.

In addition to the great faunal diversity, over 2312 hectares of old-growth forest,

including stands of red fir, lodgepole pine, Jeffrey pine, mountain hemlock, and whitebark

pine exist here (USDA Forest Service 1988). An extensive red fir forest, the only major

east-side occurrence of this species between the Kern Plateau and Hope Valley near Carson

Pass, makes up the majority of this old growth (Millar 1994).

Geographically, the San Joaquin Ridge is a low point along the Sierra Nevada

crest. Here, storms spill over from the west bringing higher precipitation to this part of the

1 eastern Sierra than to any other east-side location. This topography combined with recent, extensively destructive volcanic activity have created a migration situation for flora and fauna unparalleled anywhere on the east slope of the Sierra Nevada. These unique characteristics make this area an attractive subject for the study of floristic migration and phytogeography.

Purpose

The purpose of this study has several facets. The first is to enumerate the vascular plant taxa present in the San Joaquin Roadless Area and evaluate the floristic affinities of the taxa to place the flora in a geographic context. To this end, I will describe the historic and present climate, geology, physiography, and vegetation of the area and how these things may be affecting the present species composition and the flora's phytogeographic affinities.

I then wish to show quantitatively how the San Joaquin Roadless Area compares to other California mountain areas in its floristic composition and phytogeographic affinities.

A more local question will be to investigate quantitatively whether the flora of the area has a greater affinity to the west side of the Sierra Nevada than to other eastside floras.

To achieve these goals, I have proposed the following questions.

1. Is flora of the San Joaquin Roadless Area centered relative to the mountain ranges of

California? I expect the San Joaquin Roadless Area flora to be floristically most similar

to those from close surrounding areas, and less like those from distant mountains.

2. Are the percentages of floristic elements of geographic affinity in my flora more similar

to those of the Sierra as a whole or to those of other mountain ranges? 3

3. Due to the environmental and vegetational variation within the San Joaquin Roadless

Area, I have stratified this flora into montane, subalpine, and alpine units. Do these

elevational zone sub-floras to show different patterns of affinity?

4. How does the alpine flora of my area compare to the entire Sierran alpine? This analysis

should show how the alpine flora has developed since its disturbance compared to the

relatively undisturbed alpine flora of the Sierra as a whole.

My purpose is not to create a classification of California mountain floras according to either their species composition or phytogeographic affinities, nor to describe the floristic regions of the West. These tasks have been done by several biogeographers both in the

Great Basin and the western United States (Billings 1978, Charlet 1991, Hadley 1987,

Harper et al. 1978, Holmgren 1972, McLaughlin 1986, 1989, 1992; Pavlick 1988).

Instead, I will show how the San Joaquin Roadless Area, whose flora has been severely impacted by Holocene volcanic activity, compares to other California mountain floras and determine where the majority of its taxa have migrated from since the disturbances.

My flora is unusual in its extent of subalpine area, which is large in comparison to subalpine areas covered in other California mountain floras. I hope that this floristic study will contribute to the general knowledge of the phytogeography of California mountain ranges, and the central Sierra in particular. I also hope that the flora of the San Joaquin

Roadless Area will provide another level of precision to future quantitative studies of the mountain floras of the West, and that it will be used as a resource by both land managers and researchers. 4

The Study Area

Location and Boundaries

The San Joaquin Roadless Area is centrally located on the east side of the Sierra

Nevada at 37°42 Ν latitude and 119°02'W longitude (Figure 1). The top of San Joaquin

Ridge marks the western boundary, and a line connecting the high points of San Joaquin

Mountain, June Mountain ridge, and Peak 2987 m (more recently called 'North Point") constitutes the northern boundary. The southern boundary is the Forest Service road 3S89, and the eastern boundary is marked by Forest Service roads 3S22 and 3S89 south of

Deadman Creek and the end of Glass Flow Road (2S79) in the north. Deadman Creek

Road (3S26) extends into the study area (Figure 2).

Although the road 3S22 cuts directly through Crater Flat at the eastern edge of the study area, I included it in the survey because of its geologic and botanical interest. Crater

Flat is a low lying, seasonally wet pumice flat meadow. It is one of the lowest elevation features within the study area. The area is threatened by OHV's that stray from the road, and I felt it essential to document the plant species here.

Physiography

The study area covers about 44 km2. It spans 1136 m of elevation from 2400 m at the lowest road boundary to 3535.7 m at the top of San Joaquin Mountain. Canyons, cliffs, creeks, meadows with extensive stringers, peaks, and ridges dominate the landscape in a mix of montane and subalpine forests and woodlands topped with a small area of alpine vegetation. The underlying parent material is a mixture of volcanic, metasedimentary including limestone, and granitic rocks.

Glass Creek and Deadman Creek are the two main watercourses in the area. Each has several tributaries that drain the canyons below San Joaquin Ridge and on White Wing 5

Mountain. Smaller seasonal streams also feed these tributaries when snowmelt runoff is high.

There are five prominent peaks in the area: San Joaquin Mountain 3535.7 m, Two-

Teats 3460 m, White Wing Mountain 3051 m, June Mountain 3083 m, and Peak 2987 m

("North Point"), as well as several less prominent high points and craters, domes and flows. San Joaquin Mountain is the highest peak on the 35.7 km section of the central

Sierra crest between Blacktop peak (3902 m) on the Koip crest, and Silver Peak (3621 m) on the Silver Divide. Mammoth Mountain, which is separate from the Sierra crest, is not included in the study area.

Due to sag caused by Tertiary volcanic activity, San Joaquin Ridge, especially in the vicinity of Mammoth Pass and Deadman Pass, is the lowest point in the Sierra crest with a mean elevation 500 m lower than that of adjacent regions (Carpenter 1991, Curry

1971). Four perpendicular north-east trending spur ridges jut out from the east side of the ridge. They are separated by steep canyons and bowls that now serve as the headwaters to the several streams that flow through the study area and eventually into the Owens River.

Despite its relatively low average elevation of 3100 m, San Joaquin Ridge is a very sharply defined section of the Sierra crest. The top of the ridge itself is little more than a few meters wide in places before it drops off steeply on both sides. To the southwest it drops into the canyon of the middle fork of the San Joaquin River 600 m below. To the northeast, majestic cliffs show clear evidence of the fault block formation of the eastern escarpment that began during the late Tertiary period (Huber 1981).

Two major faults, the Fern Lake and the Hartly Springs, run parallel to the base of the escarpment south and east from just north of June Mountain to the northern edge of

Long Valley caldera. It is along these faults that the area east of the ridge broke from the rising Sierran block about 3 m.y. ago. Since then, downward displacement along these 6 faults has been about 1100 m. This break created an atypical offset or embayment within the Sierran escarpment (Bailey 1989, Huber 1981). When viewed on a map, the crest takes a distinct turn to the west before heading northward again (Figures 2 and 3).

There are six well defined meadow systems within the study area: Glass Creek

Meadows - Upper and Lower, Crater Flat, Minaret Meadow, Deadman Pass Hidden

Meadow, Deadman Canyon Alpine Meadow. The three smaller meadows lack names, so I have named them according to nearby landmarks. In addition to these well defined meadows, there are numerous meadowy areas near streams and seeps.

Glass Creek Meadow is the largest meadow system, at just under two km long. It consists of two distinct meadow areas, the upper and the lower, separated by a section of forest. Each shows distinctly different vegetation patterns. The upper has more woody plants and is wetter, while the lower is a drier grassy meadow. Several meadow stringers extend into the surrounding forest and create obscure boundaries for the meadows.

Crater Flat is a sunken area along a fault associated with the Ingo Craters volcanic chain. It is a long, shallow depression that contains a meadow on dry pumice. It receives snowmelt and runoff from a small seasonal stream that saturates the lowest part of the area in the early growing season.

Minaret Meadow is located on an unnamed stream directly east and 140 m below the Minaret Summit parking lot. Most of the meadow is dominated by willows, although there is a drier, more open section dominated by grasses, herbs, and sedges.

Deadman Pass Hidden Meadow is located at the western edge of the red fir forest at the bottom of the ridge protruding east off the north side of Deadman Pass. This meadow is completely hidden by steep rock cliffs on three sides and dense forest on the east side. It is dominated by willows, but different from the other willow dominated meadows, the very 7 wet ground between willow clumps is covered with a spongy bed of mosses through which cold-loving herbs protrude.

The Deadman Canyon Alpine Meadow system is located just below the late lying snowbanks that exist 300 m below the top of San Joaquin Ridge just south of Two Teats.

This canyon is the largest headwaters area for Deadman Creek. As the snow melts off the ridge and bowls below, it emerges in streams and seeps which feed these willow- dominated meadows. These are the coldest meadows in the study area. Ironically, the underlying bedrock is limestone, but there is so much water here that the meadow is acidic enough for a large population of ericacious plants.

Elevational Zones

Since the study area encompasses elevations from the montane to the alpine, I have divided it into elevational zones throughout the thesis to better describe various characteristics of the area and their affects on the plants.

The montane zone, defined as the upper limit of most of the red fir and Jeffrey pine in the area (Rundel et al. 1988), spans 200 m in elevation from the lower boundary of the study area at 2400 m up to 2600 m. Crater Flat, much of the red fir forest, parts of the major creeks, and the recent volcanic domes and flows are in this zone.

The subalpine zone, where mountain hemlock, whitebark pine, and limber pine are commonly found (Rundel et al. 1988), spans 400 m from 2600 m at the base of the Sierra escarpment cliff and the east end of Glass Creek Meadow up to the last upright trees at

3000 m. This elevation zone is the central core of the study area. It is highly diverse in its parent material, topography, habitats, and plant composition compared to the other two zones. 8

Using Major and Taylor's (1977) definition of alpine, the limit of upright trees, the alpine zone occupies areas above 3000 m. The alpine zone contains a large amount of vertical relief, including San Joaquin Ridge, the upper parts of the 300 to 400 m cliffs below the ridge, the upper parts of the canyons between cliff sections, and the summit areas of White Wing, Two Teats, and San Joaquin Mountain. 9

Figure 1. An outline map of California and the Ingo National Forest showing the location of the study area with major highways and towns adapted from USDA Forest Service (1995). 10

Figure 2. Topographic map of the study area showing major landmarks, scale = 1:24,000 x 40% (MaptechTM, Inc., 1997) -~- = meadow. 11

Geology

The central eastern Sierra has received extensive attention by geologists because of recent volcanic activity in and around the Long Valley caldera. A small portion of the northwest corner of the caldera is included within the study area, as well as several of the very recently active domes and craters of the Ingo Craters Chain.

Bailey (1989) has mapped the geology of the caldera and the surrounding areas that include the Ingo Craters chain and the study area. He has also provided a summary of the volcanism, tectonic structure, and glaciation of the area. Additional summaries of the recent volcanics in the area include Bailey et al. (1976), Hill (1975), Millar and Woolfenden

(1999), Miller (1985), Rinehart and Huber (1965), and Wood (1977a, 1977b, 1984).

Other studies of Long Valley caldera and Mono Craters volcanics and structure are reported in The Long Valley Symposium - Journal of Geophysical Research February 10, 1976 and

Sieh and Bursik (1986) for Mono Craters.

More recently, a large volume of research has begun due to signs of increased volcanic activity in the area: earthquake swarms, a dome-like uplift within the caldera, and carbon dioxide gas that is killing forests on Mammoth Mountain. (Hill et al. 1997, 1998;

Miller et al. 1982, Priest et al. 1998, Soret' et al. 1996).

Huber (1981) has provided a detailed history of the late Cenozoic uplift and tilt of the central Sierra, and Norris (1990) provides a good general account of the geology of the range. A brief summary and description of the geologic, volcanic and glacial features of the study area follows.

Six general types of substrates occur within the study area: Paleozoic metasedimentary rock, Mesozoic granites, Pliocene and Holocene volcanics, Pleistocene glacial deposits, and Holocene colluvium and alluvium (Bailey 1989). 12

The Paleozoic metasedimentary rock occurs along San Joaquin Ridge just below the crest. It includes a small band of limestone that outcrops in the main canyon of Deadman

Creek from about 2900 m to 3040 m. This is some of the oldest rocks in the Sierra.

The Cretaceous granodiorite pluton is exposed from White Wing Mountain east and north to June Lake, including North Point (Peak 2987 m) (Bailey 1989). Small outcrops of

Jurassic and Triassic granitic rocks are visible on the east flanks Deadman Pass.

The earliest volcanism of the region began about 3.6 m.y. ago, about the time the uplift of the central Sierra began to accelerate (Huber 1981). This is the basalt of Deadman

Pass which covered the entire San Joaquin Ridge. Today it outcrops in various places along the ridge.

San Joaquin Mountain, Two Teats, and June Mountain are quartz latite volcanoes that erupted shortly after the Deadman Pass basalt about 3.0 to 2.5 m.y. ago (Bailey 1989).

White Wing Mountain also has two Pliocene volcanic vents on its northwest end. In addition to the quartz latite flows, these eruptions also deposited a mixed quartz latite tuff and tuff breccia on areas surrounding the main vents, mostly the ridge tops and in the upper parts of the large canyons. These particular types of quartz latite are unique to the San

Joaquin Ridge area in the high Sierra Nevada (Bailey 1989, and Curry 1997 Pers. comm.).

During the Pleistocene volcanic period, several vents along the southern end of the

Fern Lake fault extruded small flows of andesite, basalt, and quartz latite. These rocks are exposed in various locations along the base of the southern half of San Joaquin Ridge.

In summary, these older rocks are exposed in several areas, creating a diversity of parent materials for development of various soil chemistries. The majority of the study area, however, is covered by much younger volcanic rocks, glacial till, colluvium, and alluvium (Figure 3). 13

Figure 3. Geologic map showing the major rock types of the study area, adapted from Bailey (1989). The large dotted line indicates the western boundary of the Long Valley Caldera. 14

Recent Volcanics

Today, most of the study area is covered with Holocene pumice tephra. This pumice blankets most of the lower elevations of the area, most of White Wing Mountain,

North Point and June Mountain. Meadows and canyon bottoms where water has been a

major factor also contain large amounts of pumice that has been reworked into colluvium and alluvium. Many of the older rocks in the area are visible because of the quick erosion of the pumice by wind and water. At Obsidian Dome and the Glass Creek and South

Deadman Flows, new land in the form of large masses of obsidian and pumice, was created by ryolite that extruded after the pyroclastic pumice eruptions.

The eruption of the Long Valley caldera, 730,000 years ago, heralded the beginning of this recent volcanic unrest in and around the area. Over the past 40,000 years, rising

magma in the area has resulted in several dozen eruptions that have formed the Ingo-Mono

Volcanic Chain.

Many of these eruptions have been in very recent times. All 21 of the Ingo craters

and domes are Holocene in age. Eruptions in the Ingo Craters Chain, beginning about 6000

years ago, have had the most effect on the area, although tephra from the Mono Craters has

also been found here (Sieh and Bursik 1986, Wood 1977b). Since about 3,000 years ago,

eruptions have occurred cyclically every 250 to 700 years.

Using radiocarbon dating procedures on charcoal logs in the ash layers, ring counts

on downed but unburned trees, obsidian-hydration rind dating, stratigraphic relations, and

sediment cores in Glass Creek Meadow, researchers have been able to construct a record

back 4,000 years B.P. dating the many different eruptions and tephra layers (Bailey 1989,

Millar and Woolfenden in press). Meadow cores show three clear layers of pyroclastic ash

deposits at 2,070 years B.P, 1670 years B.P., and 1550 years B.P. Charcoal was also 15 present in these layers indicating fires that occurred with the eruptions (Millar and

Woolfenden in press).

The most recent eruptions within and along the eastern boundary of the study area occurred 900 to 640 years ago, between ca. AD 1100 and 1360 (Millar and Woolfenden in press). Oddly, the ash layer from the last eruption at the Glass Creek vent 640 years ago

(AD 1360) is missing from the sediment cores in Glass Creek Meadow even though it is present on the slopes surrounding the meadow. Tree ring evidence shows tree death caused by tephra burial supporting the AD 1360 date from four different sample sites: San Joaquin

Ridge, White Wing Mountain, Glass Creek Forest, and North Point.

The most recent set of eruptions, located at the South Deadman, Obsidian Flow, and Glass Creek vents, discharged a total volume of 0.8 km3 of magma. Much of this magma was in the form of pumice tephra and pyroclastic flows that blanketed up to 9,000 km2 of the surrounding and downwind area (Miller 1985). These deposits traveled more than 190 km and have been found as far away as Sequoia National Park to the south and

Yosemite to the north (Miller 1985, Wood 1977b). In areas close to the vents the deposits are 10 to 25 m thick. A large part of the study area is covered with a pumice tephra blanket that is at least one meter thick (Bailey 1989). Other areas within the study area are covered with thinner tephra deposits.

These pyroclastic eruptions were quickly followed by phreatic steam explosions at the Glass Creek, and Ingo Crater Lakes, and north-east Mammoth Mountain vents. These threw more hot volcanic debris onto the surrounding area. Several of the vents then extruded rhyolite flows that are now obsidian domes.

In summary, due to the nature of these most recent eruptions, it is highly unlikely that any plants in the area survived. Miller (1985) reports that the Glass Creek tephra (AD

1360), which was the largest of the recent eruptions, killed the forest to the west of the 16

Glass Creek vent. Even if a portion of the study area's plants were not in the direct path of the most destructive pyroclastic flows, they were buried, still standing, under up to several meters of pumice tephra or burned in associated fires (Millar 1999 Pers. comm., Millar and

Woolfenden in press, Miller 1985, Wood 1977a and 1977b). Thus, the current flora in the study area has been able to grow and develop for only about 640 years since its last fiery burial. Rinehart and Huber (1965) found that pine and fir trees growing in the Ingo Craters have up to 400 growth rings, indicating a re-establishment of the forest beginning within about 200 years.

It is unclear whether some plants may have survived in a few small refugia in the lee of the tephra blasts where they were protected from burial on the west sides of the west summits of White Wing Mountain (Millar 1999 Pers. comm.) and possibly elsewhere.

There is much work to be done on the dating and mapping of individual tephra blasts that affected the area as relatively little is known.

Glacial History

There is very little exposed evidence of glaciation on and east of San Joaquin Ridge as much of the evidence of its passing has been buried by the more recent volcanics (Curry

1971, Blackwelder 1931).

Curry (1968, 1971) shows small valley glaciers of Wisconsin age, associated with the Tioga/Tenaya glaciation, in the area just below San Joaquin Mountain and Two Teats and in the two north-east facing canyons south of these peaks. Some tongues of undivided

Pleistocene glacial deposits can be found exposed below San Joaquin Mountain (Bailey

1989). At the north end of San Joaquin Ridge, a small area of Pleistocene Tahoe till runs

along the base of the ridge and covers a bench area separating the Glass Creek and 17

Deadman Creek watersheds. Today, a rock glacier formed during Holocene glacials remains in the cirque below Two Teats (Hill 1975).

Soils

Knapp et al. (1 979) and Seney (1995) describe the majority of the soils within the study area as pumice or slope wash deposits derived from pumice. Seney (1995) defined the substrate textures as gravely sand, sand, coarse sand, loamy coarse sand, loamy sand, and fine sand with less than 4% clay. These pumice soils are porous and unconsolidated with a very low water holding capacity. Very little organic matter is present. Nutrients are limited to the surface horizon.

Young age, cold temperatures, dry climate and high erosion, have all contributed to the depauperate nature of most of the soils. Only along streams and in seep, spring, and meadow areas with moisture do soils have noticeable organic matter.

Small areas throughout the study area have soils derived from granitic and metamorphic rock including some limestone (Figure 3). These soils are generally thin and rocky on slopes and in drier areas, but wetter areas such as the alpine meadows in the top of Deadman Creek Canyon support deeper mineral soils with organic material. There are also areas of bare rock and cliffs where plants survive in cracks and on ledges.

Due to the nature of pumiceous soils, water leaves the root zone quickly and there is limited water available to plants, especially during the growing season when there is little precipitation. However, when water is present, as in meadows, etc. the porosity allows pumice soils to have more available moisture than mineral soils derived from granitic and metamorphic rocks (Seney 1995). This available moisture does decrease as the depth of the pumice substrate increases. 18

Knapp et al. (1979) found that water holding capacity, depth of pumice substrate, elevation and aspect, rather than differences in nutrient availability, had the greatest affect on tree species distribution within the area. Lodgepole pine is the most capable of surviving in the deep, super well-drained pumice; whereas, red fir, western white pine, mountain hemlock, and Jeffrey pine are more common in areas with a thin coat of pumice or on rock outcrops. There are no forest trees at all where the pumice is deepest, as in Crater Flat where the tephra blanket is at least 10 m deep. I observed a few young lodgepoles invading parts of Crater Flat, but they are unhealthy and covered with snow fungus.

In Glass Creek Meadow soil pits, Millar and Wolfenden (1999) found that the pyroclastic ash is shallower to the west and deeper to the east. This corresponds to a noted increase in wet meadow plant taxa in the western portions of the meadow. The eastern part of the meadow dries out much more quickly, as evidenced by the many patches of dry meadow vegetation on the east end (Millar 1996 Pers. comm., Millar and Woolfenden in press).

Climate

Regional

The central Sierra is characterized by a montane-Mediterranean climate (Carpenter

1991). In winter, snowfall is generated by storms riding on anticyclonic winds from the

Gulf of Alaska. More than 50% of the precipitation falls between January and March with the maximum snow depth occurring in mid March (Carpenter 1991).

In the spring, subarctic air disrupts the storm cycle and brings strong winds, some extreme in velocity, up to over 25 m per second in a very short horizontal distance (<100 m) (USDA Forest Service 1995). Low pressure areas that develop in the Great Basin produce eastside northerly winds that can exceed 45 m per second. 19

By the end of April, greater than 50% of precipitation occurs as rain. Snowmelt usually begins at the end of April, but substantial amounts remain well into July, especially in shaded areas. In eleven of the past thirty years, Mammoth Mountain Ski Area has been able to remain open into July (and once until mid August) without the aid of man-made snow (Mammoth Mountain Ski Area 1999).

In the summer, the Pacific high pressure area relocates off the coast of California and blocks moisture from the north creating a perennial drought situation. This drought is only locally and sporadically relieved by thundershower activity generated by monsoonal moisture blown from the south. If the monsoons are particularly strong, rain can fall continuously for several days in a row, but even with this tropical storm moisture, less than

3% of annual precipitation falls in the summer (Carpenter 1991).

The beginning of autumn can bring more rain from tropical storms, but by the end of September, the first snowfall has often already dusted the higher peaks.

Local

Generally, precipitation decreases as you move eastward away from the crest due to the rainshadow effect. The exception is here, in and near the study area, where the average snowfall at Mammoth Pass (1270 cm) corresponds to the heaviest snowfalls recorded on the west slope at Huntington Lake in the San Joaquin drainage (Carpenter 1991).

The Mammoth Gap, a gap in the crest created by the low elevations of Mammoth,

Minaret, and Deadman Passes, combined with the alignment of the San Joaquin river drainage, allows winter storms to spill over the crest and dump their water on the east side.

The forests extending several miles east of the crest suggests this increased eastside moisture. Precipitation does still decrease as distance from the Sierra crest increases, but not as quickly as in other locations on the east side (Figure 4) and (Table 1). 20

The length of the growing season varies according to location within the study area.

Generally, it begins in May and extends to the first frost in September or October, but in severe years or cold spots at high elevations it may not begin until July and then end in late

August.

There are two distinct climatic zones within the montane portion of the study area

(USDA Forest Service 1995). One is the lower warmer, drier, northeast end of the study area that includes Obsidian Dome, Glass Creek, and the lower slopes of White Wing. The other is the upper cooler, more moist, south and west end of the study area that includes the forests surrounding Crater Flat. The drier part is much farther from the Sierra crest and is blocked by White Wing, thus receiving less precipitation than the second which tucks right up against the escarpment directly below Deadman Pass. This second area gets large amounts of snow in the winter.

Precipitation and snow depth are highest in the subalpine zone. Avalanches occur regularly in several locations below the ridges. An impressive one occurred in March of

1995, a year of exceedingly deep snowpack (Kattelmann 1996). This 9 to 12 m deep, 366 m wide wall of snow ripped 5.6 km down Deadman Creek Canyon. It uprooted 90 m of old-growth red fir forest before coming to a stop and depositing debris deep in the forest below. Some of the trees were several hundred years old and about two meters in diameter.

They were piled like spilled toothpicks at the base of the avalanche.

Steep northeast-facing canyons below the ridge are cold air corridors that hold snow late into the growing season. In contrast, the wide open, more gentle canyon that holds Glass Creek Meadow, though under deep snow in winter, is blooming yellow with buttercups long before the steeper canyons are even accessible. White Wing, June

Mountain, and North Point are even more exposed to sun and wind and are free of snow earlier in the season, except in north- and east-facing shaded areas. 21

Within the alpine, snow accumulation varies from little on west-facing ridges to very deep in the more protected, wind-loaded east slopes. The zone is characterized by cool summers with high near ground temperatures, wide diurnal temperature fluctuations, severe winds, drought, cold, and the possibility of summer freezes (Major and Taylor

1977). 22

Figure 4. Sierra Neveda precipitation, monthly means from east and west slope stations for 58 years (Carpenter 1991). 23

Weather Data

There are no weather recording stations within the study area, and those surrounding the area are not consistent in the types of data they report. I have put information together from these nearby locations to provide a reasonably accurate picture of the precipitation, snow depth, and temperatures experienced in the study area (Table 1).

The two closest weather stations are Mammoth at 2530 m and Gem Pass at 3277 m.

These two stations are at comparable elevations to the study area and are located just to the north and south. They provide measures of snow depth and water content for the area.

Mammoth Mountain Ski Area records actual monthly snowfall.

Knapp et al. (1979) used 1964 data from Ellerly Lake, Gem Lake, and Convict

Lake stations to estimate average January and July temperatures and precipitation at Minaret

Summit. They found average lows to be around -4°C and average highs around 16°C with extreme lows of -29°C and extreme highs of 30 to 32°C. They estimated precipitation to be about 76 cm with 80% falling as snow.

Average annual precipitation from 1941 to 1970 at Gem Lake is 52 cm and at

Ellerly Lake is 57 cm (California Department of Water Resources 1980) (Figure 5). Gem

Lake is at 2744 m in elevation, 533 m below Gem Pass.

More recent data show the 1969 to 1999 average snow depth at Gem Pass to be 207 cm with a water content of 79 cm. April first average snow depth is 81 cm (CA Dept. of

Water Resources 1999). These data are most comparable to the climate along San Joaquin

Ridge and the canyons below it (Table 1).

At Mammoth, the 1990 to 1999 average snow depth is 136 cm with a water content of 47 cm. April first average snow depth is 51 cm (CA Dept. of Water Resources 1999).

These data are most comparable to the red fir forest and Glass Creek Meadow. (Table 1). 24

Snowfall totals for Mammoth Mountain average 900 cm per year since 1969 (Mammoth

Mountain Ski Area 1999) (Figure 6).

The closest station that currently records temperatures is Lee Vining, 48 km to the north. Lee Vining is at 2069 m in elevation at the base of the Sierra escarpment. Average temperatures in Lee Vining from 1988 to 1998 range from -1°C in January 1020°C in July

(Western Regional Climate Center 1999). These temperatures are probably warmer than most areas in the study area which are at higher elevations. To illustrate the effect of the

Mammoth Gap, average annual precipitation in Lee Vining is 38 cm, less than half of what the study area receives (Table 1). Table l. Summary of average annual weather data for stations near the San Joaquin Roadless Area. Minaret Summit is situated on the southwestern boundary of the study area, and is probably most representative of the average climate of the area. n/a = not available. (a = California Department of Water Resources 1999, b = California Department of Water Resources 1980, c = Knapp et al. 1979, d = Western Regional Climate Center 1999)

Station Elevation Snow depth (cm) Water content of Precipitation (cm) Temperatures (°C) Snowpack (cm) Avg. High Avg. Low Gem Pass a 3277 m 207 79 n/a n/a n/a n/a n/a n/a n/a Ellerly Lake b 2896 m 57 Minaret Summit c 2796 m n/a n/a 76 16 -4 n/a n/a n/a n/a Gem Lake b 2744 m 52 n/a n/a n/a a 47 Mammoth 2530m 136n/a n/a Lee Vining d 2069m 38 20 -1 25 26

Figure 5. Preciptiation and temperature at Gem Lake (elevation 2744 m) based on a 60 year mean (Carpenter 1991). 27

Figure 6. Snowfall totals for Mammoth Mountain Ski Area 1969-1999 (Mammoth Mountain Ski Area 1999). 28

Human Impacts

Throughout the last century, grazing, timber harvest, and recreation have been the major sources of human impacts on the study area. Small scale mining has occurred sporadically in the area since before the turn of the century, but its impact was localized and minimal (Knapp et al. 1979). The area has not burned in the last 125 years (Millar and

Woolfenden in press).

Sheep grazing has had the greatest impact on the vegetation in the northern part of the study area. It was especially heavy from the mid 19th century until 1950 when regulations were put on the number of days sheep could be in the area. Until recently, permits allowed 1800 sheep to travel through the grazing allotment from July 1 to August

31. This rate typically meant that the sheep would be in a meadow for a total of six to ten days (USDA Forest Service 1995).

Two locations long used as bedding areas, one near a spring on the north side of lower Glass Creek Meadow and the other on the bench between the Glass and Deadman watersheds, are dominated by the native but weedy Chenopodium atrovirens. A small population of the alien Bromus tectorum also exists on the south-facing upper slope of the bench.

Interestingly, pollen cores from Glass Creek Meadow have shown no significant differences in vegetation composition between the pre-grazing period and the last 150 years when grazing occurred regularly (Millar and Woolfenden in press). It seems that the effects of grazing are visible, but they are not on a scale large enough to be detected by pollen analysis procedures.

Corn lilies (Veratrum californicum), indicators of heavy grazing, dominate parts of upper Glass Creek Meadow, and there is noticeable stream channel incision on the part of

Glass Creek that flows through the meadow. There are also four alien plant species 29

(Polygonum arenastrum, Taraxicum officinale, Rorippa nasturtium-aquaticum, and

Veronica serpyllίfolίa ssp. humίfusa) widespread throughout the meadow. Iris missouriensis, indicated as a noxious weed by Hickman (1993), is present in the meadow, but is not a dominant as in other heavily grazed meadows on the east side. Today grazing is not permitted in Glass Creek Meadow (USDA Forest Service 1997, Millar and Woolfenden in press).

Some timber harvest has occurred within the study area, but not to any great extent.

The red fir near the end of upper Deadman Road was logged in 1962 and 1963 and lower elevation forest on the south and east slopes of White Wing were logged some time between 1982 and 1992. (USDA Forest Service 1995, Leven 1992). In 1986, the Forest

Service decided that no logging would occur in the red fir belt at the base of San Joaquin

Ridge (USDA Forest Service 1995).

Recreation is the most widespread human activity in the study area, though historically it has had relatively low impact as few people come into the roadless area. Most recreation use is hunting, fishing, camping, hiking, and cross-country skiing. There are only a couple of designated hiking trails and no developed campgrounds so recreation use is minimal and dispersed, except where motorcyclists have illegally ridden off road in

Crater Flat, along Glass and upper Deadman Creeks, and on the summit and slopes of

White Wing (Knapp et al. 1979, Leven 1992, personal observations 1994-1997).

Developed recreation, however, is the greatest threat to the flora of the study area.

In the late 1970's, the roadless area was left out of the John Muir Wilderness in order to maintain the site's potential as a downhill ski area that would connect the Mammoth and

June Mountain resorts (USDA Forest Service 1995). The impact from this type of development could be severe. As things stand today, the Forest Service has determined that the building of ski lifts within the study area is not compatible with the desired condition 30 for that location, but it has not completely eliminated the possibility of heli skiing in the area and an expansion of the June Mountain ski area just outside the boundaries of the study area (Friends of the Ingo 1996, USDA Forest Service 1995, Sally Miller 1999 Pers. comm.).

The greatest threat is the plan to build mountain bike trails throughout the roadless area, specifically along the top of San Joaquin Ridge. This subalpine and alpine perennial habitat is sensitive to disturbance and also home to one of the two rare plant species found in the site. Unfortunately, I have witnessed mountain bikers lack of respect for designated trails and seen many off trail pumice plants crushed by bike wheels. This type of developed recreation has not been determined incompatible with the desired condition (Friends of the

Ingo 1996). Developed bike trails and heli skiing are two activities that, once sanctioned by the Forest Service and implemented, would close the possibility of this roadless area becoming part of the adjacent John Muir Wilderness.

Research and Botanical Collecting History

The San Joaquin Roadless Area and its surroundings have been the focus of many scientific studies and recorded observations (Burke 1982; Bailey et al. 1976; Bailey 1989;

Curry 1966, 1968, 1971; Giuliani 1990; Hill et al. 1997, 1998; Huber 1981; Ivey 1979;

Kilbourne and Anderson 1981; Leven 1992; Lipshie 1976; Millar 1994; Miller et al. 1982;

Miller 1985; Rinehart and Huber 1965; Smith 1976; USDA Forest Service 1988; Wood

1984, 1977a, 1977b), but there has been very little botanical collecting in the area.

Most of the study has occurred within the last ten years as a result of the Mammoth to June Ecosystem Analysis pilot project for the Sierra Nevada Ecosystem Project. This study encompassed many topics ranging from the geology, forest stand structure, and wildlife to air or water quality and social issues (USDA Forest Service 1995, Millar 1996, 31

USDA Forest Service 1997). Following is a summary of the botanical work done in and around the study area.

Millar and Woolfenden (1999) used dendrochronology, wood anatomy, forest stand and structure analysis, and pollen analysis to research the historic conditions of Glass

Creek Meadow and surrounding forests over the past 4000 years. Carpenter (1991) did a red fir and lodgepole pine seedling establishment study at Deadman Creek. Potter (1994)

included 13 plots from the eastside red fir forest in his ecological classification of Sierran upper montane forests. Knapp et al. (1979) did an ecological study of Minaret Summit in order to determine the extent of biotic interchange across the crest. This study included a

list of plants occurring in the southern end of my study area and was the only focused botanical collecting done in the area.

Botanical collecting has occurred just outside both the southern and northern

boundaries of the study area. Ann Howald conducted a survey of the vegetation and flora of Mammoth Mountain for the Mammoth ski area (Howald 1983), and Mark Bagley conducted a sensitive plant survey on June Mountain for the June ski area (Bagley 1988).

Just to the west, the National Park Service is in the process of compiling a checklist of

plants in Devils Postpile National Monument (U S Department of the Interior 1996).

Vegetation

Paleovegetation

The history of the vegetation of California and the Sierra Nevada in particular has

been covered by many authors (Axelrod 1976, Chabot and Billings 1972, Major and

Bamberg 1963 and 1967, Millar 1996, Raven and Axelrod 1978, Stebbins and Major

1965, Woolfenden 1996). I will describe here only the pertinent points that relate to the

recent development of the San Joaquin Roadless Area flora. 32

Recent dendrochronoligical work on White Wing Mountain and pollen cores in

Glass Creek Meadow show strong evidence that, especially within the last 2000 years, the vegetation was severely affected by repeated volcanic disturbance and climate change

(Millar and Woolfenden in press).

Α "ghost forest" on the summit of White Wing Mountain has provided several clues into the recent past of the study area. This forest is made up of hundreds of dead trees, all lying with their tops facing away from the Glass Creek vent. These trees were apparently snapped from their bases by the force of the last eruption (640 years ago) but were spared from fire. Because of the cold, dry climate, the wood is well preserved.

Tree ring counts, wood anatomy studies and carbon 14 dating have shown that this forest was a mixed forest of Jeffrey pine, lodgepole pine, whitebark pine, western white pine, and sugar pine living ca. 650 to 1100 years ago. The dates of the living forest coincide with the Medieval Warm Period (900 -1300 years ago), and the death date coincides to the Glass Creek eruption and the second long drought of the warm period

(Millar and Woolfenden in press, Woolfenden 1996).

The identification of sugar pine as part of this 900 year old forest is significant.

Today sugar pine does not occur on the east side of the Sierra except near Lake Tahoe 200 km to the north, and the closest sugar pine to White Wing is 30 km to the west and 750 m lower in elevation (Millar and Woolfenden in press). It is possible that during the Medieval

Warm Period there may have been additional taxa growing here on the east side with the sugar pine, that today are restricted to the west side of the Sierra.

Soil pits in the red fir forest have revealed better developed soil profiles beneath the current layer of pumice tephra (Knapp et al. 1979), showing further indications of different conditions in the recent past. This could be due to a combination of events including a 33 longer period between volcanic disturbances and a warmer climate with more summer moisture.

Two factors have since prevented this rich conifer forest from regenerating on the summit of White Wing, which today is barren pumice except for a few krummholz whitebark pines. The thick layer of tephra deposited by the blast created a much more difficult situation for tree regeneration. In addition, coinciding with this last eruption, was the beginning of the Little Ice Age (ca. AD 1400) which brought a cooler climate, drier summers, and a shorter growing season. Pollen cores from Glass Creek Meadow show an increase in cold-loving plants at 450 years ago, also coinciding with the Little Ice Age

(Millar and Woolfenden in press, Woolfenden 1996).

As a result of these disturbances over the past several thousand years, the study area flora has had only the last 600 years or so to develop free of volcanic obliteration and major climate change events. Millar and Woolfenden (1999) note that the vegetation of

Glass Creek Meadow is surprisingly high in biomass and species diversity given the fact that it has been so frequently disturbed, and has had such little time to recover.

Current Vegetation

The vegetation of the Sierra has been described by many authors, of which Smiley

(1921) and Sharsmith (1940) are most classic. I use Sawyer and Keeler-Wolfs (1995) statewide classification to outline the vegetation types found within the study area. This system defines a set of major vegetation types called "series" based on dominance and characteristic species. They are equivalent to alliances of the Nature Conservancy's

National Vegetation Classification (Grossman et al. 1994).

Series or alliances are considered to be collections of associations, vegetation units defined on the present characteristic species. Series broken down into associations are used 34 to describe kind of vegetation with consistent species composition in the central Sierra by many authors including Benedict (1983), Burke (1982), Halpern (1986), Pemble (1970),

Ratliff (1982, 1985), Major and Taylor (1977), and Taylor (1979, 1984). These contain many descriptions for non-woody types, especially at the higher elevations.

Groups of associations that share similar environmental conditions but have not yet been categorized as series are described as habitats by Sawyer and Keeler-Wolf (1995).

Habitats are used to describe many high elevation areas that lack the extensive plot-based sampling needed to classify them into series.

In my description of the study area's vegetation I replace the term "series" with a descriptive structural term such as forest, woodland, shrubland, and chaparral. Α shrubland is an area dominated by shrubs; whereas, a chaparral is a shrubland dominated by evergreen shrubs with sclerophyllous leaves. In addition, I use the following terms: stand and montane chaparral. Α stand is an area in which the vegetation is uniform and can be classified into a series. Montane chaparrals differ in species composition from low elevation chaparrals. They often occur well into subalpine elevations.

I use my observations and field notes in order to provide a general picture at the middle scale series, association, and habitat levels as defined in Sawyer and Keeler-Wolf

(1995). I also refer to Woolfenden et al. (1995), who provide an overview of the forest and shrub types at the series-level based on plot data measuring cover, life form, and size classes. Since that study is most concerned with forest management, it is not as detailed in regards to non-forested types.

I divide the study area into its elevational bands (montane, subalpine, and alpine) to more clearly present the vegetation. These bands are defined in the Elevational Zones section of the study area description. 35

Montane Zone

The montane zone vegetation consists of forests, wet and dry meadows, and streamside shrub habitats. The lower, drier portion of the study area is covered by a mixed forest of red fir, Jeffrey pine, western white pine, and lodgepole pine. This forest can be considered a higher elevation form of Jeffrey pine forest (Sawyer and Keeler-Wolf 1995).

Common understory plants include Purshia tridentata var. tridentate, Penstemon rostriflοrus, Lupinus argenteus and Ribes cereum. This forest occurs around the north and east base of White Wing and along Glass Creek.

A small amount of Jeffrey pine forest lacking red fir, western white pine, and lodgepole pine occurs on the south side of Obsidian Dome near the Glass Creek vent.

Several Great Basin taxa are in the understory including Cercocarpus ledifolius and

Chamaebatiaria millefolium. This forest extends higher into the subalpine zone on North

Point with Purshia tridentata var. glandulosa as a dominant understory shrub.

The most prominent vegetational feature of the upper part of the montane zone is the red fir forest (Sawyer and Keeler-Wolf 1995). The vast expanse of eastside red fir forest occurs between 2500 m and 2600 m in the pumice flats at the base of San Joaquin Ridge and extends higher in places. This area receives deep, late lying snow required to support a red fir forest (Barbour et al. 1991). The porous nature of the pumice soil allows adequate drainage as red fir does not tolerate saturated soil (Oosting and Billings 1943). The understory is sparse; Pyrola picta and Pedicularis semibarbata are often the only herbs present. Most of the herb diversity occurs along the stream courses.

This red fir forest differs in composition from the extensive pure stands of the west side in that it contains scattered trees of lodgepole pine, Jeffrey pine, western white pine, mountain hemlock, and quaking aspen (Millar 1994, USDA Forest Service 1995). Millar

(1994) mentions a note from Dr. Susan Ustin (UC Davis) that this red fir is genetically 36 unusual as certain morphological and biochemical traits align it more closely to var. shastensis.

A small area of lodgepole pine forest (Sawyer and Keeler-Wolf 1995) occurs at the eastern edge of the red fir forest around the edges of Crater Flat at 2500 m. Lodgepole pine also occurs in disturbed areas within the red fir forest (Woolfenden et al. 1995).

Wetland shrub and meadow habitats occur along streams and near springs throughout the zone and extend into the subalpine zone along stream corridors where conditions are favorable. Ratliff s (1982, 1985) Carpet clover association is found along stream edges among willow thickets. Brayshaw's (1976) descriptions of montane wetland shrub habitat apply to wet areas where Salix lemmonii is dominant. An extensive dry meadow area at Crater Flat is dominated by Carex douglasii.

Subalpine Zone

The vegetation here is a patchy mosaic of forests, meadows, and shrublands as a result of a great topographic variety.

Stands dominated by whitebark pine, mountain hemlock, lodgepole pine, or limber pine, and mixed subalpine forest apply to Sawyer and Keeler-Wolfs (1995) series. Limber pine occurs only on the upper north slopes and summit of North Point as Taylor's (1979)

Limber pine/curlleaf mountain mahogany association.

Lodgepole pine forests are most extensive on the dry pumice areas around the north base of White Wing and on moist meadow margins where the tree mixes with willows. In the dry forests understory composition is often similar, yet lower in diversity to the open pumice habitat.

Mixed subalpine forest (Sawyer and Keeler-Wolf 1995) is the dominant forest type to the west of Glass Creek Meadow and along the base of San Joaquin Ridge above the red 37 fir forest. The conifers represented here are red fir, lodgepole pine, western white pine, mountain hemlock, and whitebark pine. The understory is predominantly occupied by shrubs; however, in places it is very sparsely vegetated with herbs such as Lupinus lepidus and Poa wheeleri.

Small mountain hemlock forests occur in the coldest, somewhat protected locations, usually on north facing canyon sides and cliffs and at the base of the escarpment. Many alpine taxa make their way into the subalpine zone in this habitat. This forest is very limited. It is usually a combination of red fir and mountain hemlock (Woolfenden et al.

1995). The understory often consists of Luzula spp., Phyllodoce breweri, and Primula suffructescens.

Whitebark pine woodlands occur high on canyon walls and lower, more protected ridge tops below San Joaquin ridge and its associated spur ridges. Here upright trees grade into krummholz as conditions change toward an alpine environment. Eriogonum microthecum, Valeriana californica, and Arnica nevadensis, as well as many alpine plants are found in the sparse understory.

Shrublands consist of areas dominated by short, shrub-high aspen, sagebrush, and several different montane chaparral species. Short aspen covers large areas on the south slope of June Mountain and in the Deadman Creek Canyons, especially in avalanche paths.

Aspen groves of taller trees also occur in isolated wet areas in the red fir or mixed subalpine forests, and along watercourses. The understory is sparsely dominated by Symphoricarpos

rοtundifοlius, Lupinus argenteus, Elymus elymoides, and Αchnatherum occidentale.

There are several montane chaparrals inhabiting non-forested steep and/or rocky slopes below the low sagebrush shrubland or in forest openings. At the higher elevations

Tobacco brush or Greenleaf manzanita chaparrals are the most extensive. Bitterbrush dominates exposed mid-elevation ridges. At lower elevations and in forest openings on the 38 southeast sides of White Wing Bush chinquapin chaparral and Huckleberry oak chaparral can be found (Sawyer and Keeler-Wolf 1995).

Areas dominated by sagebrush. Low sagebrush shrubland (Artemisia arbuscula) (Sawyer and Keeler-Wolf 1995) covers high elevation, south facing, steep slopes and mid-elevation, exposed ridges. A distinct break between this shrubland and the larger shrubby montane chaparrals is striking, especially on the upper slopes of June

Mountain, the southwest peaks of White Wing, and the bench between the Glass and

Deadman watersheds. On the upper west part of this bench is a unique assemblage of low shrubs dominated by low sagebrush and an uncommon, high elevation, pink color morph of Castilleja linearifolia (Howald 1983). Here, the snow melts and the plants bloom before anywhere else at comparable elevations.

Big sagebrush shrubland (Artemisia tridentata ssp. vaseyana) (Sawyer and Keeler-

Wolf 1995) occurs throughout the subalpine and montane zones and as an understory shrub in the forests and in montane chaparrals. It is a dominant in the dry areas of the avalanche path in the bottom of the middle canyon of Deadman Creek and on the bench between the Glass and Deadman watersheds. Artemisia tridentate ssp. tridentate does not grow within the boundaries of the study area, but it dominates lower elevations to the east.

In areas where the tephra is deepest and extremely well-drained, forests and chapparals give way to a assemblage of low, deeply tap-rooted Great Basin desert perennials dominated by Parry rabbitbrush (Sawyer and Keeler-Wolf 1995). This open pumice shrubland dominates White Wing Mountain in the subalpine and the dry portions of

Crater Flat in the montane. The rare Mono Lake lupine (Lupinus duranii) grows in this shrubland at the base of White Wing on the northeast side.

Within the open pumice habitat, subtle differences in the chemical make-up of the pumice and the texture and size of soil grains appear to cause sharp changes in vegetation 39 pattern. It is clearly demonstrated in the Lupinus duranii population near the northeast base of White Wing. A visible line between fine, gray pumice and coarse, reddish/orange pumice runs perpendicular to the slope. There are no changes in slope angle or aspect, but on the gray, fine-grained pumice L. duranii is the dominant plant with about 50% cover.

Just meters away, on the reddish/orange coarse-grained pumice there are very few plants of any species and no Lupinus duranii. A study on the autecology of this rare plant would greatly benefit from plots in this area.

The subalpine zone has extensive riparian and meadow areas. Most of the riparian areas are thickly lined with willows, primarily Salix geyeriana, S. jepsonii, S. lemmonii, and S. orestera, that can be assigned to Sawyer and Keeler-Wolfs (1995) subalpine wetland shrub habitat. Here, Burke's (1982) Colville ragwort-showy sedge association occurs in wetlands and along streamsides in the canyons. Major and Taylor's (1977) Sierra willow-arrowhead butterweed association is found in Glass Creek Meadow. Hidden

Meadow is covered with Taylor's (1984) Grayleaf willow-meadow onion association.

Most of the meadows and meadowy areas around springs and along riparian corridors are generally described as Sawyer and Keeler-Wolfs (1995) subalpine meadow habitat.

The vegetation of lower Glass Creek Meadow goes through three stages of dominance throughout a growing season. From snowmelt through early summer the meadow is bright yellow, dominated by two subspecies of Ranunculus alismifolius. Before the hottest part of the summer, Ratliffs (1982, 1985) Longstalk clover association dominates the meadow, turning it pink. As the summer wears on, the meadow turns white as it changes into a patchwork of Benedict's (1983) Heretic penstemon - yarrow association, sedges, and Halpern's (1986) Rough bentgrass and Tall manngrass associations. Major and Taylor's (1977) Pussypaws - heretic penstemon association occupies the drier edges of the meadow and large areas of Minaret Meadow. Benedict's 40

(1983) Many nerved sedge - yarrow association occurs in meadowy areas in the canyons, in Glass Creek Meadow, and at Crater Flat in the montane zone. Upper Glass Creek

Meadow is dominated by willows, mostly Salix geyeriana and S. lemmonii, and large patches of Delphinium polycladon and Veratrum californicum.

Alpine Zone

The alpine zone is characterized by areas of wet and dry meadows, whitebark pine krummholz, low alpine shrublands, and fell fields. Small riparian areas run through the wet meadows.

The lowest parts of the alpine are along San Joaquin Ridge near Deadman Pass, in the top of the middle canyon of Deadman Creek, and in the large bowl below San Joaquin

Mountain. The Deadman Pass area supports a subalpine upland shrub habitat that is dominated by Major and Taylor's (1977) Granite gilia-alpine goldenbush association. The top of the middle canyon of Deadman Creek supports sedge (Carex lenticularis) dominated wet meadows in which Dodecatheon alpinum, Salix arctica, and Veronica wormskjoldii are found. The bowl below San Joaquin Mountain is a large open pumice-like area dominated by Taylor's (1984) Nuttal sandwort and Parry rush-vagus buckwheat associations.

Higher on the ridges whitebark pine krummholz becomes dominant with a sparse understory of Eriogonum lobbii, Lupinus lepidus var. lobbii, and Penstemon davidsonii.

Here, the rare Arabis pinzlae is a distinct member of the understory on south-facing slopes.

On the very tops of these ridges, the krummholz gives way to Major and Taylor's (1977)

Granite gilia-alpine goldenbush association.

The highest 200 m of the study area near San Joaquin peak differs greatly from the surrounding highlands, both in parent material and plant composition. Small dry sedge meadows near the peak are dominated by Carex tahoensis, C. praticola, and C. albonigra. 41

Fell fields are dominated by Major and Taylor's (1977) Watson spikemoss-roundleaved buckwheat and Alpine pussypaws-heretic penstemon associations, first described by

Pemble (1970). The most distinct assemblage in the summit zone, however, is a group of alpine cushion plants dominated by Lupinus lepidus var. ramosus, a lupine found nowhere else in the study area. Within both the subalpine and alpine zones are extensive rock outcrops. These areas support their own list of rock-loving plants such as Heuchera micrantha var. erubescens, Melica stricta, Oxyria dignya, and Penstemon newberryi. 42

Figure 7. Glass Creek Meadow and San Joaquin Ridge from the east, July 10, 1995.

Figure 8. Glass Creek Meadow and White Wing Mountain from the summit of June Mountain, July 16, 1996. Figure 9. San Joaquin Ridge from the southern edge of Deadman Pass looking north, August 18, 1995.

Figure 10. The east-side red fir forest beneath San Joaquin Ridge, looking south from White Wing Mountain, July 13, 1995. METHODS

Field Methods

I used the U.S. Forest Service delineation of the San Joaquin Roadless Area

(USDA Forest Seryice 1995) to determine the boundaries of my study area. I excluded

Yost Meadow and the Hartley area, north and east of June Mountain ridge line, which are outside the Glass and Deadman watersheds (Figure 2).

I obtained collecting permits from Ingo National Forest botanist Kathleen Nelson and wildlife biologist Richard Perloff along with watch lists of potential rare and sensitive plants in the area. Bagley's (1988) sensitive plant survey of the nearby June Mountain Ski

Area provided field identification characteristics of rare and sensitive plants that I might encounter. I also consulted floras from nearby and adjoining areas for lists of plants I might expect to find in the study area (Bagley 1988, Howald 1983, Taylor 1981).

During the summer and fall seasons of 1994, 1995, and 1996, one day in 1997 and two days in 1999, 1 conducted 65 days of field work in which I collected vascular plants in the area. I collected and identified 1236 specimens that are deposited at the Humboldt State

University Herbarium (HSC).

Despite the difficulty of thoroughly surveying such a large and topographically diverse area, I made an effort to cover the area by collecting plants in each vegetation type.

Special emphasis was given to meadows and riparian areas, the alpine zone, and open pumice areas; as well as different rock types, slope angles and aspects. I intensified surveys in the wet areas where the flora was more diverse as opposed to the relatively homogeneous forest understory. It was therefore not necessary for me to hike through every km2 of land (Table 2). In each vegetation type I passed through, I collected specimens of all species in flower or fruit except for the widespread, already collected and known species. For these I made notes.

44 45

By spreading the collecting over several years, I was able to visit the vegetation types at different times during the growing season. In addition, each year was very different in terms of snowpack depth and the timing of peak bloom. In 1994 the snow melted early and the buttercups in Glass Creek Meadow were in full bloom by June 4. 1 collected many late blooming species that year.

The 1995 season came after a very heavy winter. On June 30, the snow had not completely melted from Glass Creek Meadow, and the exposed buttercups were still in bud. Peak bloom did not happen until mid August. During this season, I followed the snow melt and was able to collect the earliest blooming plants. I also collected in September and October for later plants. This year saw a great change in parts of the area as several major avalanches opened previously forested areas.

The 1996 year was also wet and cold with a snow storm on June 30. In contrast to

1995, however, many early plants were blooming by this time, and some were frost damaged. This was also the wettest summer on the east side in several years, as a result of frequent thunderstorms. During the 1996 season, I was able to collect many early, middle, and late blooming plants. l also found several new plants in areas where I had already collected the previous years.

In 1997 I collected in the limestone area at peak bloom, as the plants had past bloom by the time I found it the year before. In 1999,1 returned to the location where I found the rare Arabis pinzlae to better document its population. Throughout my field work, l took many photographs to record vegetation and topographic characteristics of the area (Figures

7, 8, 9, and 10).

By the end of the 1996 season, I was finding few or no new species so I ended

collecting, except for the couple of confirmation hikes in later years. There are still areas I would like to collect more thoroughly, especially in the more difficult to reach areas high in 46

the canyons and on the limestone outcrop which I found at the end of my field work. I was

also unable to collect on and below the many cliff faces due to the danger of falling rock.

Specimen Identification Methods

I identified my specimens using keys in the Jepson Manual (Hickman 1993) corrected according to Schmid (1999), with additional reference to Munz (1959, 1968) and

Cronquist et al. (1972-1994). 1 used Weeden (1986) in the field and Mason (1957),

Hitchcock and Chase (1971), Abrams (1940-1960), and Hurd et al. (1998) to help identify some specimens. I also consulted the HSU Herbarium, and gained professional confirmation on the rare plants (J. Morefield - Arabis and T. Sholars - Lupinus). I have filed National Diversity Data Base (NDDB) field survey forms for the rare plants in the study area. 47

Table 2. Locations of important collecting areas. I collected in large radii around these central points.

Collecting Area Elevation Townsihp & Range Latitude Longitude Meadow near Deadman Group Camp 2420 m Τ.2S. R.27Ε. Sec 32 37°43'55"N 119°00'59W Jeffrey pine forest near the south side of 2424 m Τ.2S. R.27Ε. Sec.32 37°44'10"Ν 119°00'42"W Glass Creek Flow Crater Flat 2498m Τ.3S. R.27Ε. Sec.8 37°41'22N 119°01'12"W Along streams in red fir forest below 2498 m Τ.3S. R.27Ε. Sec.7 37°42'09"N 119°02'13"W Deadman Pass Lower Glass Creek 2517 m Τ.2S. R.27Ε. Sec.19 37°44'45"Ν 119°01'24"W Mixed forest near end of the third right 2530 m Τ.3S. R.27Ε. Sec.20 37°40'27"N 119°01'15"W off Dry Creek Road South side of White Wing 2549 m Τ.3S. R.27Ε. Sec.6 37°43'21 "N 119°01'40"W Mixed Jeffrey pine forest near end of 2554 m Τ.2S. R.27Ε. Sec.30 37°44'45"Ν 119°01'33"W Glass Flow Road East slope of Glass Creek Flow 2560 m Τ.2S. R.27Ε. Sec.20 37°44'35"N 119°01'20"W Red fir/Jeffrey pine forest near end of FS 2562 m Τ.2S. R.27Ε. Sec.31 37°43'50"N 119°01'30"W Road 2S50, left spur End of Deadman Creek Road 2582 m Τ.3S. R.26Ε. Sec.8 37°42'50"Ν 119°03'16"W Rocky pumice mound along trail tο 2582 m Τ.2S. R.27Ε. Sec.30 37°44'38"Ν 119°01'40"W Glass Creek Meadow, Lupinus duranii location Hill in red fir forest just west of Crater 2613 m Τ.3S. R.27Ε. Sec.18 37°41'07"Ν 119°02'01 "W Flat Red fir forest along the closed end of 2619 m Τ.3S. R.26Ε. Sec.2 37°42'43"Ν 119°03'43"W Deadman Road Mixed Jeffrey pine forest east of Glass 2631 m Τ.2S. R.26Ε. Sec.35 37°44'40"Ν 119°01'57"W Creek Meadow South facing, steep pumice slopes 2677 m Τ.2S. R.27Ε. Sec.30 37°44'43"Ν 119°02'10"W northeast of White Wing Pumice flats northwest of White Wing 2683 m Τ.2S. R.27Ε. Sec.30 37°44'34N 119°02'32"W Avalanche zone in bottom of the middle 2687 m Τ.3S. R.26Ε. Sec.2 37°42'55"Ν 119°04'22"W canyon of Deadman Creek Red fir/mountain hemlock forest in the 2726 m T.3S. R.26Ε. Sec.11 37°4221 "Ν 119°04'15"W south canyon of Deadman Creek Minaret Meadow 2731 m Τ.3S. R.26Ε. Sec.25 37°3927"Ν 119°0252"W Hidden Meadow 2732m Τ.3S. R.26Ε. Sec.12 37°41'45"N 119°03'35"W Glass Creek Meadow, lower 2743 m Τ.2S. R.26Ε. Sec.36 37°44'08"Ν l 19°03'15"W Upper Glass Creek Meadow and 2743 m Τ.2S. R.26Ε. Sec.35 37°4353"Ν 119°03'46"W stringers Shepherd Camp Spring, north side of 2755 m Τ.2S. R.26Ε. Sec.36 37°44'17"N 119°03'20"W Glass Creek Meadow Northeast slopes of White Wing 2777 m Τ.2S. R.27Ε. Sec.30 37°44'23"Ν 119°02'02"W 48

Table 2. Important collecting areas, continued.

Collecting Area Elevation Townsihp & Range Latitude Longitude South facing slopes below Peak 2865m 2802m Τ.3S. R.26Ε. Sec.1 37°43'08"Ν 119°03'31 "W Bench between Glass and Deadman 2822 m Τ.3S. R.26Ε. Sec.2 37°43'15"N 119°04'36W watersheds Southeast slopes of White Wing 2841 m Τ.2S. R.27Ε. Sec.31 37°43'51"N 119°02'04"W Bowl below Deadman Pass 2846m Τ.3S. R.27Ε. Sec.13 37°41'26"Ν 119°03'46"W South Canyon of Deadman Creek 2856 m Τ.3S. R.26Ε. Sec.11 37°42'01 "Ν 119°04'25"W 37°42'37"N Limestone outcrop in the middle canyon 2900 m Τ.3S. R.26Ε. Sec.3 119°04'59W of Deadman Creek South slopes of the south canyon of 2908 m T.3S. R.26Ε. Sec.11 37°42'26"N 119°04'30"W Deadman Creek South slope of June Mountain 2924 m Τ.2S. R.26Ε. Sec.35 37°4420"Ν 119°03'49W Deadman Pass, ridgetop and cliffs below 2982 m Τ.3S. R.26Ε. Sec.14 37°41' 19"Ν 119°04' 11 "W Subalpine forest along north tributary to 3000 m Τ.3S. R.26Ε. Sec.2 37°43'15"N 119°04'54W Deadman Creek Metamorphic spur ridge off San Joaquin 3020 m Τ.3S. R.26Ε. Sec.13 37°41'44"N 119°04'13W Ridge near Deadman Pass Alpine meadows in the middle canyon of 3020 m T.3S. R.26Ε. Sec.3 37°42'32"N 119°05'14"W Deadman Creek North slopes of the south canyon of 3030 m Τ.3S. R.26Ε. Sec.11 37°42'08"N 119°04'48"W Deadman Creek Summit areas of White Wing 3042m Τ.3S. R.26Ε. Sec.1 37°43'31 "Ν 119°02'42"W Summit of June Mountain 3047 m Τ.2S. R.26Ε. Sec.26 37°44'23"Ν 119°04'09"W South ridge of the south canyon of 3056 m Τ.3S. R.26Ε. Sec.11 37°42'21"N 119°04'50"W Deadman Creek Upper north tributary to Deadman Creek 3123 m Τ.3S. R.26Ε. Sec.4 37°43'07"N 119°0518"W Spur ridge to the east of San Joaquin 3189 m Τ.3S. R.26Ε. Sec.2 37°43'23"N 119°05'23"W Mountain San Joaquin bowl 3228 m Τ.3S. R.26Ε. Sec.4 37°43'09"N 119°05'37"W Spur ridge east of Two Teats, Arabis 3229 m Τ.3S. R.26Ε. Sec.3 37°43'03"N 119°05'35W pinzlae location San Joaquin Ridge, near Peak 3290m 3290m Τ.3S. R.26Ε. Sec.11 37°42'09 Ν 119°05'10"W Two Teats 3413m Τ.3S. R.26Ε. Sec.3 37°42'43"Ν 119°05'57"W San Joaquin Peak 3536 m Τ.3S. R.26Ε. Sec.4 37°43'07"Ν 119°06'14"W 49

Analysis Methods

Floristic studies can be used to study the affinities of a taxon to a particular region as well as to study the geographic affinities of complete floras (Stott 1981). Studies of phytogeographic affinities of both taxa and floras has been subjective until the recent use of

Q- and R-mode factor analyses (McLaughlin and Bowers 1990, McLaughlin 1992). Here the phytogeographic affinities of taxa were determined subjectively, but the quantitative techniques of cluster analysis and ordination were used to determine the floristic affinities of my flora.

Phytogeographic Affinities of the Taxa

To determine the floristic makeup of a flora, the phytogeographic affinities of each taxon need to be determined. Several factors must be taken into account. First is the center of origin (Stott 1981, Morefield 1992), which can be determined by the following criteria:

(1) the center of taxanomic diversity (Daubenmire 1978, Gleason and Cronquist 1964,

Raven and Axelrod 1978, Stebbins 1982), (2) the center of abundance (Cain 1944, Wulff

1950), and (3) the location of primitive forms (Raven and Axelrod 1978, Stebbins and

Major 1965, Stebbins 1982). Cain (1944) provides descriptions of these and more criteria along with their assumptions.

Phytogeographic affinities can also be evaluated by considering genetic relationships, especially based on ploidy levels (Daubenmire 1978, Raven and Axelrod

1978, Stebbins 1982, Stott 1981); similarities in ecological conditions where groups of taxa grow (Billings 1976, Bliss 1985, Chabot and Billings 1972, Daubenmire 1978); and evolutionary history based on fossils (Cain 1944, Daubenmire 1978, Raven and Axelrod

1978, Reveal 1979, Stebbins 1982, Stott 1981, Wulff 1950). 50

This task requires detailed knowledge of many specialties (Charlet 1991). I am especially indebted to long time scholar of Sierran floristics, Dr. G. Ledyard Stebbins, who assisted me in these determinations. As the original data from his 1982 paper were missing, he graciously reviewed my flora and personally assigned each taxon to its phytogeographic element according to his previous work (Stebbins 1997 Pers. comm.). Table 3 defines

Stebbins' six phytogeographic elements.

I then assigned each taxon in the other 12 floras to its phytogeographic element using its range, its center of abundance based on species descriptions, and its center of taxanomic diversity using Hickman (1993), Munz (1959, 1968), Cronquist et al. (1972-

1994), and Hitchcock and Chase (1971). In addition I considered the judgements of

Chabot and Billings (1972), Major and Taylor (1977), Morefield et al. (1988), Polunin

(1959), Raven and Axelrod (1978, 1985), Reveal (1979), Sharsmith (1940), Stebbins

(1982), and Taylor (1977).

Ideally, published maps, monographs, and herbarium specimens should be used to determine a taxon's phytogeographic affinities (Morefield 1992), but the size of this project precluded my referencing the primary information for each of the 2,296 taxa. 51

Table 3. Phytogeographic elements as defined by Stebbins (1982, 1997 Pers. comm).

Old Cordilleran The taxon's probable center of origin and close relatives occur in Rocky Mountains and/or Cascades.

Circumboreal The taxon's probable center of origin and close relatives occur throughout the Holarctic area, including low elevations in the Pacific Northwest.

Lowland California The taxon's probable center of origin and close relatives occur in the California Floristic Province of cis-montane California and southwest Oregon, including the Sierra Nevada.

Great Basin The taxon's probable center of origin and close relatives occur in the Great Basin/Desert regions, mostly within and above the pinyon/juniper zone east of the Sierra Nevada.

Northern Hemisphere, The taxon is widespread in the temperate regions of eastern no particular region North America. These taxa were originally categorized as Circumboreal by Stebbins (1982) and are combined with that group in the affinities analyses for direct comparison to his work.

Alien The taxon is not native to California. 52

Comparison of My Flora with Other Floras

I selected 13 high elevation floras in California based on the following criteria: (1) availability and relative completeness of the species list, (2) location of the units within or on the border of California, and (3) coverage of montane through high alpine habitats.

Tuolumne Meadows and Vicinity and the Hall Research Natural Area, which cover only subalpine and alpine areas, were used since they are near mine. Beyond these, I selected floras that are located in the cardinal directions from my area (Table 4). After my initial analysis I eliminated Devils Postpile as the list lacked many grasses, sedges, and willows.

Since these floras were compiled at different times throughout this century, I needed to stabilize nomenclature to that of Hickman (1993). I maintained subspecies and variety ranks consistently throughout. In many cases, I was able to assign missing subspecies names in older floras using the modern synonymy. I eliminated hybrids and taxa reported as a questionable occurrence within a flora.

A difficult problem in comparing floras (Operational Geographic Units or OGUs,

Crovello 1981) is how different a flora's area is from the others (Beals 1984, Goodall

1973). Recent studies in Europe overcome size/area problems by dividing an area into equal sized plots, surveying and mapping species, and ordinating the plots. Some of these regions are quite large, such as the national grid of Britain (Hill 1991), though usually much smaller in size and scope (Andersson and Weimark 1996, Bolognini and Velluti

1995, Heikkinen 1998, Nimis 1997). This approach was not practical for me, since knowledge of occurrence of a species is not available in subfloras of smaller plot sizes than the study area.

In order to make the OGUs comparable, I excluded taxa from the flora that occur below the montane elevation. My low elevation cut levels were 1829 m in the south and

1524 m in the north. In addition, I performed analyses on subsets: 1) floras with similar 53 numbers of taxa to mine, 2) floras near to mine, and 3) floras located in the Sierra Nevada

(Table 5).

Phytogeographic Affinities of the Floras

In the first analysis I followed a common method (Chabot and Billings 1971,

Krantz 1994, Lavin 1983, Major and Bamberg 1963, Major and Taylor 1977, Mitchell

1973, Potter 1983, Sharsmith 1940, Smiley 1921, Stebbins 1982) of comparing percentages of each phytogeographic element present in each flora with a simple chart.

In addition, I stratified the taxa into those that grow in the montane only, the montane and subalpine, the subalpine only, the subalpine and alpine, the alpine only, and in all three zones to investigate the details in my flora as it compares to Stebbins findings about the entire Sierra Nevada.

I further follow other studies (Billings 1978, Estrada-Loera 1991, Hadley 1987,

Harper et al. 1978, Lavin 1983, Peterson 1986) by comparing the floras using similarity coefficients. I then use them to perform cluster analyses (Hadley 1987, Nimis 1997, Taylor

1977), and ordinations of the floras (Charlet 1991, Kirkpatrick 1982, Taylor 1977). I used

Sorensen's coefficient for presence-absence data (Sorensen 1948) to perform cluster analyses (Aldenderfer and Blashfield 1984, Kent and Coker 1992) and Bray-Curtis ordinations (Bray and Curtis 1957) on the PC ORD computer program (McCune and

Mefford 1995).

Similarity Coefficient. The Sorensen's coefficient is preferred for this kind of study (Cottam et al. 1973, Taylor 1977). It looks at the similarity between samples, giving weight to species in common rather than to species absent. It is relatively robust with samples with high 13-diversity (range of compositional difference between the floras)

(Goodall 1973, Kent and Coker 1992, Whittaker and Gauch 1973). 54

Cluster Analysis. A cluster analysis (Aldenderfer and Blashfield 1984, Kent and Coker 1992) presents the flora relationships as a dendrogram, a pictorial representation of the relative similarities among floras. This method is often used to create classifications

(Aldenderfer and Blashfield 1984), but in my case I used the cluster analysis to help answer my questions.

I used a hierarchical agglomerative method with the complete linkage, also called the farthest neighbor, algorithm (Sokal and Michener 1958). This algorithm clusters based on similarity to all floras in the group, rather than to only one (Aldenderfer and Blashfield

1984). This method produced groups that made the most geographic sense (Aldenderfer and Blashfield 1984).

Ordination. Ordination techniques are commonly used to study vegetation patterns related to environmental controls (Kent and Coker 1992). I chose to use a Bray-

Curtis ordination with variance-regression endpoint selection with the city-block measure to calculate residual distances (Beals 1984). This technique assumes linear species distributions, causing some distortion with high 13-diversity (Charlet 1991, Whittaker and

Gauch 1973), but it provides the smallest amount of distortion when using presence- absence data (Cottam et al. 1973, Kent and Coker 1992, Whittaker and Gauch 1973).

Since it is susceptible to very different floras, obscuring the relationships among the floras

(Beals 1984, Cottam et al. 1973), I also performed analyses on subsets of the 13 flora set

(Table 5).

The Bray-Curtis ordination is an indirect gradient analysis, based on floristic data independent of possible controlling environmental factors (Kent and Coker 1992). To determine if environmental variables had any correlation to the relationships of the floras, I overlaid a secondary matrix with these variables onto the species by flora ordination

(Appendix B). Table 4. California mountain floras used in the statistical analyses. *Area size codes: 1 = smaller than mine; 2 = same size (approx.) as mine; 3 = larger than mine; 4 = a regional flora, much larger than mine.

Name Reference Elevation Size* Latitude Longitude Sαπαnaquinadlessea,nounty,n California H. Constantine-Shull 2400 - 3536 m 2 37°39' - 37°45'Ν 119°00' - 119°03'W (2000) Harvey Monroe Hall Research Natural Area, Mono E. Knapp (1993) 2927 - 3838 m 1 37°55' - 37°59'Ν 119° 16' - 119° 18'W County, California White Mountains of Mono and lnyo counties, J. Morefield et al. (1988) 2378 - 4343 m 4 37° 10'- 37°57'Ν 117°58' - 118°25'W California and Esmeralda County, Nevada Upper Walker River Watershed - Sierra Nevada M. Lavin (1983) 2 378 - 3793 m 3 38°03' - 38°40'Ν 119° 15' - 1 19°38'W portion, Mono County, California Sweetwater Mountains, Mono County, California M. Lavin (1983), K. Hunter 2378 - 3559m 3 38°16' - 38°36'Ν 119°14' - 119°25'W and R. Johnson (1983) Bodie Hills, Mono County, California and Mineral T. Messick (1982) 2439 - 3121 m 2 38°06' - 38°23'Ν 119°14' - 119°46'W County, Nevada Tuolumne Meadows and vicinity, Tuolumne County, J. T. Howell (1944) 2348 - 3838 m 2 37°50' - 37°57'Ν 120°13' - 120°24'W California Lassen Volcanic National Park, Shasta, Lassen, and G. Gillette et al. (1961) rev. 1585 - 3188 m 3 40°25' - 40°35'Ν 121°15' - 121°35'W Plumas counties, California by V. Oswald et al. (1995) Trinity Alps, Trinity County, California W. Ferlatte (1974) 1524 - 2745 m 4 40°47' - 41 °01'N l22°47' - 123°16'W Sαn Bernardino Mountains, San Bernardino County, T. Krantz (1994) 1829 - 3505 m 4 34°00' - 34°22'Ν 116°37' - 117° 16'W California Sequoia and Kings Canyon National Parks, Tulare and J. Rockwell and S. 2134 - 4419 m 36°16' - 37°14'N 118°25' - 119°02'W Fresno counties, California Stocking (1969), U. S. Geological Survey (1998) Desolation Wilderness, EIElrado County, California B. Potter (1983) 2256 - 3044 m 2 38°50' - 39°04'Ν 120°06' - 120°15'W Mammoth Mountain, Mono County, California A. Howald (1983) 2439 - 3370 m 1 37°40' - 37°42'Ν 119°01' - 1 19°02W 5 5 56

Table 5. Subsets of the 13 floras used in the analyses. Numbers in parenthesis indicate number of taxa.

SIMILAR-SIZED FLORAS NEARBY FLORAS SIERRAN FLORAS

San Joaquin Roadless Area (443) San Joaquin Roadless Area San Joaquin Roadless Area White Mountains (516) Hall Research Natural Area Hall Research Natural Area Bodie Hills (329) Upper Walker River Upper Walker River Tuolumne Meadows (475) Sweetwater Mountains Tuolumne Meadows Trinity Alps (573) Bodie Hills Sequoia and Kings Canyon Desolation Wilderness (532) Tuolumne Meadows National Parks Mammoth Mountain (287) Mammoth Mountain Desolation Wilderness Mammoth Mountain RESULTS

Floristic Analysis

The vascular plant flora of the San Joaquin Roadless Area consists of 446 taxa, including 48 families, 176 genera, and 416 species. Asteraceae with 73 taxa, Poaceae with

50 taxa, and Cyperaceae with 32 taxa comprise 35% of the flora. Carex is the largest genus with 30 taxa. Other large genera are Arabis and Eriogonum (12); Lupinus and Epilobium

(11); Poa (10); and Artemisia, Erigeron, and Juncus (8).

Rare and Uncommon Taxa

There are two rare (California Native Plant Society - CNPS List 1B; Skinner and

Pavlick 1994) taxa in the area. I found a population of at least 100 plants of Arabis pinzlae at an elevation of 3200 m to 3280 m on the southeast-facing side of the east trending spur ridge below Two Teats. Individuals are widely scattered among whitebark pine krummholz just below the ridge top. This is the first report of this plant in the Sierra Nevada, and it is a western range extension for the species that was previously only found on Boundary Peak in the White Mountains (Morefield 1994).

This taxon appears to be a close relative Arabis platysperma var. platysperma, which also grows in the area. Morefield and Taylor (1990) found plants intermediate in plant robustness, and mature leaf and stem pubescence between the taxa. This intermediacy has spawned a question as to their taxonomic status. In the San Joaquin population, all of the plants had the typical A. pinzlae hair branching and density and leaf and fruit size, but there were two plants that had mature fruits 0.5 mm longer than expected for A. pinzlae.

Differences between the Sierran and the White Mountains populations of A. pinzlae occur in terms of habitat as well. In the White Mountains, it occurs on north- and east-

57 58 facing granitic scree slopes. In my area it grows on south- and east-facing volcanic scree slopes.

Α large population of at least 30,000 plants of Lupinus duranii exists along the trail to Glass Creek Meadow and on the south side of Glass Creek at the north-east foot of

White Wing Mountain. It is a dominant in the Parry rabbitbrush scrub and also an understory component of the Jeffrey pine forest. The soil is a fine-grained pumice sand with larger pumice gravel. This work verifies the presence of Lupinus duranii populations west of U.S. 395. The greatest threat to this population is illegal mountain- and motor-bike riding in the area. Foot traffic along the trail to Glass Creek Meadow is another impact.

Carex praticola and Chaenactis douglasii var. alpina, both found in the alpine zone near San Joaquin Peak, and Carex petasata, found in a seep on the north side of the south canyon of Deadman Creek, are listed as rare in California but more common elsewhere

(CLAPS List 2). Astragalus kentrophyta var. danaus, found in the alpine zone near San

Joaquin Peak, and Eleocharis parvula, found in Glass Creek Meadow, are on the CLAPS watch list (List 4).

Three taxa are considered uncommon (not very likely to be encountered, but not rare enough to be designated as sensitive or included in the CLAPS inventory; Hickman

1993). Two Eriogonum taxa are fairly common in my area. Eriogonum rosense grows on the alpine ridges below San Joaquin Peak and below White Wing Mountain with Lupinus duranii. Eriogonum microthecum var. alpinum grows on the bench between the Glass and

Deadman watersheds, on metamorphic rock in the alpine, and below White Wing Mountain with L. duranii. Carex tahoensis grows in the alpine on San Joaquin ridge and below the peak. 59

Range Extensions

My work extends the ranges of 3 taxa and documents previously unknown populations of 3 taxa growing here at the edge of their ranges. All four cardinal directions are represented.

My specimen of Artemisia michauxiana is the first report of this species in the

Sierra Nevada, except for one undocumented sighting in Glacier Canyon in Yosemite

(Howell 1944). In California, 11 specimens are from the White and Ingo mountains (Cal

Flora 1999, Hickman 1993) and one from the Marble Mountains (Howell 1939, Smith and

Sawyer 1987). Outside California the plant is widely distributed in the Rocky Mountains.

My report of Artemisia ludoviciana ssp. candicans is the first specimen in Mono

County (Cal Flora 1999), and extends its range to the south. It typically occurs from

Donner Pass in the Sierra Nevada and the Modoc Plateau, north to Washington, Montana, and Utah (Hickman 1993).

Delphinium gracilentum is the only species in my area considered an exclusively west-side taxon in range. Maps and listings from the Cal Flora Database show it occurring in moist meadow habitats from 0 to 3048 m all along the west side of the central Sierra.

The only location where this plant has been found on the east side of the Sierra is on

Mammoth Mountain (Howald 1983). I found an additional small population deep in the willows in Minaret Meadow. These populations extend the range of this taxon to the east.

Mine is the first report of Penstemon heterodoxus var. cephalophorus in Mono

County. It is typically found in the southern Sierra Nevada at high elevations in the vicinity of Sequoia and Kings Canyon National Parks in Tulare and Fresno counties.

Although Pinus flexilis is known from several locations in Mono County, the population on top of North Point (Peak 2987 m) is an additional and previously unknown 60 occurrence of this typically southern Sierra Nevada tree. The northernmost occurrence in the Sierra is at Twin Lakes near Bridgeport (Cal Flora 1999).

My collections of Purshia tridentata var. glandulosa confirm a previously known northern range extension of this variety (USDA Forest Service 1995). This variety occurs primarily in the desert ranges to the south of the study area.

Alien Taxa

As described in the human impacts section of the introduction, five alien and three weedy native taxa grow in my area.

Other Noteworthy Taxa

Small populations of 9 taxa are restricted to San Joaquin Peak:

Antennaria media Cryptantha sobolifera Astragalus kentrophyta var. danaus Lupinus lepidus var. ramosus Carex albonigra Phlox condensata Carex praticola Ivesia lycopodioides ssp. lycopodioides Chaenactis douglasii var. alpina

Several other taxa occur only as a single occurrence in the study area. Antennaria dimorpha occurs on pumice soils at Crater Flat growing at its upper elevational limit

(Hickman 1993). One small population of Cirsium scariosum occurs in a small, steep, intermittent stream course that feeds into the north canyon of Deadman Creek. A few plants of Descuriana incana occur on one, very exposed basalt outcrop. Dodecatheon alpinum,

Salix arctica, and Ledum glandulosum occur in the alpine meadows in the middle canyon of

Deadman Creek. A small population of Mimulus moschatus occurs on one sandy, steep, wet slope of the middle canyon of Deadman Creek.

One small Populus balsamifera ssp. trichocarpa tree occurs in the red fir forest.

This tree is next to a now closed road that was historically used by miners and sheep 61 herders and more recently by hunters and wood gatherers. Populations of Sibbaldia procumbens and Lithophragma glabrum occur near a rocky spring area below San Joaquin bowl. One small population of Sorbus californica occurs only in riparian areas in the south canyon of Deadman Creek. Tetradymia canescens occurs only in the quartz latite rocks at the summit of June Mountain.

Noteworthy Occurrences on the Limestone

Although I did not found any rare or widely disjunct taxa on the limestone, the plant composition differed considerably from that of other rock types. Limestone occurs around

2900 m on both sides of the middle canyon of Deadman Creek either as south-facing cliffs or as small north-facing rock outcrops.

The interesting assemblage I examined is growing on the dry, north-facing outcrops. The following taxa are found on the limestone:

Abies magnίfica var. magnίfica Penstemon newberryi var. newberryi Arabis lemmonii var. depauperate Phlox diffusa Arabis platysperma var. platysperma Pinus albicaulis Arenaria kingii var. glabrescens Poa cusickii ssp. epilis Artemisia ludoviciana ssp. ludoviciana Poa secunda ssp. secunda Artemisia tridentata ssp. vaseyana Poa wheeleri Cymopteris terebinthinus var. petraeus Silene bernardiana Cystopteris fragilis Symphoricarpos rοtundifοlius var. Lupinus argenteus var. argenteus rotundifolius Monardella glauca Trisetum spicatum Pellaea breweri Valeriana calίfornίca

In addition, Swertia radiata and Castilleja applegatei ssp. pallida occur only here in the study area. Two stunted trees of Pinus jeffreyi, Angelica lineariloba (which is typical in montane chaparral), and Stephanomeria tenuifolia (which grows on pumice soils in the montane forests) occur as disjunct, high-elevation populations here. A disjunct, low- 62 elevation population of Selaginella watsonii, otherwise found in the study area only on San

Joaquin Peak, occurs here as well.

Noteworthy absences

Abies concolor occurs nearby on Mammoth Mountain and in the June Lake area but

does not occur in the study site. Deschampsia cespitosa is a common meadow grass in the

Sierra Nevada, however, I found only D. elongata. Juniperus communis occurs near June

Lake and was reported from the area (Curry 1994 Pers. comm.), but I did not locate it.

Sedum pinetorum was reported in 1913 from near Pine City, an old mining camp

near Lake Mary, just a mile from Mammoth Mountain. Dean Taylor alerted me of its

possible occurrence (1994 Pers. comm.); however, since then, CLAPS determined that this

taxon does not occur in California (Skinner and Pavlick 1994). I did not encounter any

succulent plants.

Statistical Analysis

Comparison of All Floras

The species composition of my flora is most similar to that of Tuolumne Meadows

and vicinity (56%), and least to the San Bernardino Mountains (30%). The average

similarity between all floras is 39%. The two most similar floras are the Sierran portion of

the Upper Walker River watershed and the Sweetwater Mountains (85%), while the San

Bernardino Mountains and the Hall Research Natural Area are most dissimilar (15%)

(Appendix C).

The cluster analysis (Figure 11) suggests three major groups of floras, 1) Sierran

and Westside floras, 2) Great Basin and eastside floras, and 3) floras not similar to either 63 group (Lassen National Park, Trinity Alps, San Bernardino Mountains). My flora is included with other Sierran and Westside floras, and is most similar to its geographically closest floras, and then to the other Sierran floras on the west side of the crest.

The Bray-Curtis ordination shows more or less the same pattern (Figure 12). My flora is compositionally centered, close to its geographic neighbors, but positioned on the

Great Basin side of the Sierran group. The Great Basin floras are positioned toward the far right side of the ordination. Along the first two axes, the Trinity Alps, Sequoia and Kings

National Parks, and Hall Research Natural Area are most distant based on the interpretation of these axes as flora size and location. Along axis three the San Bernardino Mountains are distant also based on location (Figure 13).

In Figures 12 and 13 the three axes account for only 29% of the variation in species composition of the 13 floras. Overlaying my environmental variables explained some of the relationships among floras. The first axis is most strongly correlated with the east-west distance from my flora (r2 = 0.639). The second axis is most strongly correlated with the number of taxa (r2 = 0.575). The third axis is most strongly correlated with the north-south distance from my flora (r2 = 0.421) (Tables 6 and 7).

The positioning of the floras in my ordination compared to their size and actual geographic locations support these findings. However, the three axes account for very little of the variation and few of the environmental variables correlations with the axes have an r-squared that exceed 0.50. This indicates that there is a large variation among the floras, and that there are additional factors contributing to this variation.

Due to high B-diversity and the small amount of variation explained by the three axes in the complete ordination, I tried several analyses on subsets of the flora set. These provide a more detailed understanding of the relationships between the floras. In these analyses, the three axes account for > 70% of the variation between the floras. 64

San Joaquin Roadless Area

Mammoth Mountain

Tuolumne Meadows

Desolation Wilderness

Sequoia/King's Canyon National Parks Hall Research Natural Area

White Mountains

Upper Walker River

Sweetwater Mountains

Bodie Hills

Lassen National Park

Trinity Alps

San Bernardino Mountains

Figure 11. Cluster analysis of the complete set of floras. 65

Figure 12. Ordination of all floras, axes one and two.

Figure 13. Ordination of all floras, axes one and three. 66

Table 6. Coefficients of determination for the correlations between ordination distances and distances in the original n-dimensional space from the analysis of all floras.

Axis R Squared increment Cumulative 1 .060 .060 2 .068 .128 3 .160 .288

Table 7. Pearson correlations of environmental variables with ordination axes from the analysis of all floras.

Axis 1 Axis 2 Axis 3 r2 r2 r2 Latitude Midpoint .132 .006 .265 Latitude Difference .350 .031 .421 Longitude Midpoint .395 .002 .165 Longitude Difference .639 .039 .206 Elevation Span .001 .484 .110 Highest Elevation .242 .219 .001 Number of Taxa .003 .575 .227 67

Comparison of Floras of Similar Size

Α cluster analysis from a subset of floras based on size maintains the groups found by the original analysis (Figure 14). My flora continues to be grouped with the Westside

Sierran floras.

In an ordination, the three axes account for 77% of the variation among the floras

(Table 8). My flora is centered together with Mammoth Mountain and Desolation

Wilderness. Except for changing Tuolumne Meadows location, it agrees with the

ordination for the complete flora set (Figure 15). 68

San Joaquin Roadless Area

Tuolumne Meadows

Desolation Wilderness

Mammoth Mountain

White Mountains

Bodie Hills

Trinity Alps

Figure 14. Cluster analysis of floras of similar size.

Figure 15. Ordination of floras of similar size. 69

Table 8. Coefficients of determination for the correlations between ordination distances and distances in the original n-dimensional space from the analysis of floras of similar size.

Axis R Squared increment Cumulative 1 .074 .074 2 .569 .643 3 .128 .770 70

Comparison of Floras Close to the San Joaquin Roadless Area

A cluster analysis (Figure 16) resulting from this subset agrees with the original analysis by grouping my flora with Sierran Westside floras, but it differs by including

Mammoth Mountain with the Great Basin and eastside group. An ordination supports the original findings by positioning Mammoth Mountain in the center (Figure 17). The three axes account for 75% of the variation among the floras. (Table 9). 71

San Joaquin Roadless Area

Tuolumne Meadows

Hall Research Natural Area

Upper Walker River

Sweetwater Mountains

Bodie Hills

Mammoth Mountain

Figure 16. Cluster analysis of floras close to the San Joaquin Roadless Area.

Figure 17. Ordination of floras close to the San Joaquin Roadless Area. 72

Table 9. Coefficients of determination for the correlations between ordination distances and distances in the original n-dimensional space from the analysis of floras close to the San Joaquin Roadless Area.

Axis R Squared increment Cumulative 1 .274 .274 2 .279 .553 3 .193 .746 73

Comparison of Sierran Floras

A cluster analysis on this subset shows my flora to be most similar to the eastside floras (Figure 18). The Westside floras are then grouped together, with the Hall Research

Natural Area separate because of its lack of montane taxa.

An ordination centers my flora with one eastside flora, Upper Walker River watershed, and one west-side flora, Desolation Wilderness. Sequoia and Kings Canyon

National Parks and the close geographical neighbors are positioned around the edges

(Figure 19). The three axes account for 74% of the variation among the floras (Table 10). 74

San Joaquin Roadless Area Mammoth Mountain

Upper Walker River Tuolumne Meadows

Desolation Wilderness Sequoia/King's Canyon National Parks Hall Research Natural Area

Figure 18. Cluster analysis of Sierran floras.

Figure 19. Ordination of Sierran floras. 75

Table 10. Coefficients determination for the correlations between ordination distances and distances in the original n-dimensional space from the analysis of Sierran floras.

Axis R Squared increment Cumulative 1 .332 .332 2 .268 .268 3 .137 .737 76

Floristic Affinities of the San Joaquin Roadless Area

The San Joaquin Roadless Area contains a full half of the Sierran flora as presented by Stebbins (1982). The phytogeographic affinities of the San Joaquin Roadless Area flora are generally consistent with those of the entire Sierran flora (Table 11, Stebbins 1982).

However, within each affinity category, the numbers differ. Old Cordilleran taxa make up the largest group (48.4%), a substantially higher percentage compared to that of the Sierra

Nevada as a whole (39%). Circumboreal and Northern Hemisphere taxa comprise the next largest group (20.9%), a low amount compared to the whole Sierra (26%). Lowland

California make up 16.1% of the flora compared to 19% for the entire Sierra. Only 13.5% of the flora originated in the Great Basin. For an eastside flora, this number is low, as the entire Sierra has 16% of its flora originating in the Great Basin.

I compared the affinities in the alpine zone of San Joaquin Roadless Area to

Stebbins' (1982) study. The proportions of affinities in my alpine parallel those of the entire Sierran alpine, except for Lowland California taxa (Table 12), where I had twice that of Stebbins. The proportions of Circumboreal taxa do not increase in my alpine as they do by 2% in the entire Sierran alpine, and those of my alpine Great Basin taxa do not decrease as they do by 1% in the entrie Sierra (Table 13). 77

Table 11. Comparison of the phytogeographic affinities of my flora with those of the entire Sierran flora (Stebbins 1982).

Flora Old Circumboreal + Lowland Great Basin Alien Total Cordilleran N. Hemisphere California

San Joaquin Roadless Area 216 (48.4%) 93 (20.9%) 72 (16.1%) 60 (13.5%) 5 (1.1%) 446 Sierra Nevada (Stebbins 1982) 342 (39%) 227 (26%) 168 (19%) 139 (16%) N/A 876

Table 12. Comparison of the phytogeographic affinities of my alpine flora with those of the entire Sierran alpine flora (Stebbins 1982).

Flora Old Circumboreal + Lowland Great Basin Alien Total Cordilleran N. Hemisphere California

San Joaquin Roadless Area Alpine 105 (54%) 36 (19%) 25 (13%) 27 (14%) 0 193 Sierra Nevada Alpine (Stebbins 1982) 103 (50%) 58 (28%) 15 (7%) 31 (15%) N/A 207

Table 13. Phytogeographic affinities of the San Joaquin Roadless Area by elevation zone.

Old Circumboreal + Lowland Great Alien Total Cordilleran N. Hemisphere California Basin Zone Montane restricted (8) 31% (3) 12% (6) 23% (9) 35% 0 26 Subalpine restricted (68) 45% (37) 25% (26) 17% (15) 10% (5) 3% 151 Alpine restricted (38) 49% (16) 20% (11) 14% (13) 17% 0 78

Occur in montane (79) 52% (27) 18% (26) 17% (19) 13% 0 151 Occur in subalpine (169) 50% (73) 22% (54) 16% (38) 11% (5) 1% 1% 339 Occur in alpine (105) 54% (36) 19% (25) 13% (27) 14% 0 193 78

Upon examining the percentages of taxa in each affinity category for all floras, I found that the affinities of each flora support the relationships uncovered by the species composition analyses. My area is geographically centered in the mountains of California, so it is expected that the percentages in each affinity category for my flora fall close to the center of the affinities reported for the other 12 floras. Figure 20 shows that my area is indeed intermediate in terms of the proportions of its flora that belong to each affinity category.

The proportions shown by the other floras also reflect their geographic distribution

(Table 14). The continuous habitat available along the Sierra/Cascade axis has allowed floras in these areas to be dominated by taxa from the Old Cordilleran and Circumboreal regions. The floras with the largest proportions of taxa from these regions (>55% and

>15% respectively) are the close geographic neighbors to my area: Hall Research Natural

Area and Tuolumne Meadows, probably because of their lack of montane taxa. The Trinity

Alps also contain a high percentage of these taxa (52.9% and 10.3%), especially considering their relatively low elevation and lack of extensive higher elevation habitats.

Floras in the southern and eastern geographic positions contain relatively low proportions of Old Cordilleran and Circumboreal plants, all < 45% and <12% respectively. The San

Bernardino Mountains and Bodie Hills floras contain the lowest percentages of these taxa.

My flora falls near the center with 48.4% Old Cordilleran and 13.5% Circumboreal plants.

The Westside Sierran and northern floras show low affinities to the Great Basin

(<10%), while the eastside and Great Basin floras contain high percentages. The Bodie

Hills and the White Mountains have the greatest affinity to the Great Basin with more than

25% of their taxa originating there. My flora, along with Mammoth Mountain and the San

Bernardino Mountains, falls in the middle with about 14% Great Basin taxa. 79

Just the opposite is true for the Lowland California derivation category. Those floras high in Great Basin taxa typically show low affinities to the California Floristic

Province. As expected, the White Mountains contain by far the lowest percentage of

Lowland California taxa (4.7%), while they are second only to the Bodie Hills for the highest percentage of Great Basin taxa. However, of the floras that occur east of the Sierra crest, my flora and Mammoth Mountain contain the highest percentage of Lowland

California taxa with 16.1% and 19.5% respectively.

The Northern Hemisphere derivation category reflects widespread taxa, many of which prefer lower elevations or are aquatics. Lassen National Park contains the highest percentage of these taxa, while the Hall Research Natural Area contains the lowest. Again, my flora contains an average proportion of these taxa.

Besides Tuolumne Meadows and Hall Research Natural Area, my flora contains the smallest proportion of alien plants. Mammoth Mountain, my area's closest geographic neighbor and a heavily impacted ski area, has the highest percentage of alien plants. Only one flora, the Hall Research Natural Area has reported zero alien plants.

A cluster analysis of the percentages in each affinity category supports the findings of the other analyses in this paper by creating two main groups, 1) Great Basin and eastside

Sierran floras and 2) Sierran and western floras (Figure 21). The only major difference is that this analysis does not group my flora and Mammoth Mountain together. Instead, it groups my flora with the Sierra and western floras, and Mammoth Mountain with the Great

Basin and eastside Sierran floras. However, it continues to show the similarity of my flora to its other close geographic neighbors, Tuolumne Meadows and Hall Research Natural

Area. 80

on. Cany s ing K

C = K

h. t u so h to t nor from d e ang arr e r a as lor F

ry. o teg ca ity n fi f a hic rap eog h g eac in

as f flor o t en c Per

0. 2

e ur ig F 81

Table 14. Number of taxa in each affinity category for California mountain floras, arranged north to south. (KC=Kings Canyon).

Flora Old Circum- Lowland Great Northern Alien Total Cordilleran boreal California Basin Hemisphere

Trinity Alps 303 59 223 116 43 19 573 Lassen National Park 360 95 157 36 87 50 785 Desolation Wilderness 271 64 101 40 41 15 532 Upper Walker River 311 '79 85 131 60 40 706 Sweetwater Mountains 286 59 70 137 48 38 638 Bodie Hills 130 24 35 107 26 7 329 Hall Reseach Natural Area 116 39 30 11 6 0 202 Tuolumne Meadows 265 69 75 32 30 4 475 San Joaquin Roadless Area 216 59 72 60 34 5 446 Mammoth Mountain 130 25 56 42 9 25 287 White Mountains 232 58 24 148 40 14 516 Sequoia/KC National Parks 401 105 210 79 67 19 881 San Bernardino Mountains 267 59 223 116 81 50 796 82

San Joaquin Roadless Area Hall Research Natural Area

Tuolumne Meadows

Lassen National Park Sequoia/King's Canyon National Parks Desolation Wilderness Trinity Alps

White Mountains

Bodie Hills

Upper Walker River

Sweetwater Mountains

Mammoth Mountain

San Bernardino Mountains

Figure 21. Cluster analysis of percentages in each affinity category. DISCUSSION

The San Joaquin Roadless Area flora is central, relative to other California

mountain floras, in its geographic location, species composition, and phytogeographic

affinities. Severe and recent volcanic disturbances within the past 1,000 years created a

new, non-vegetated landscape open to immigration and re-colonization by plants from

surrounding areas. This new landscape is characterized by conflicting extremes of wet and

dry environments created by unusually heavy east-side snowfall combined with large

expanses of quickly draining, sandy pumice substrates. These extremes have come together

here to create a moderate environment that supports a newly immigrated, centrally aligned

flora — one that is not strongly aligned to either the east or west side of the Sierra, but rather to the Sierra in general.

Floristic Affinities of the San Joaquin Roadless Area

Species Composition Analyses

The statistical analyses confirm the central nature of the San Joaquin Roadless Area

flora. Species composition comparisons to other Sierran floras show that my flora has

strong affinities to the eastside floras of Mammoth Mountain and the Upper Walker River

watershed, as well as to the Westside flora of the Desolation Wilderness (Figures 18 and

19). When other mountain areas are compared, my flora is consistently grouped with one

or more of three Westside floras: Tuolumne Meadows, Desolation Wilderness, and/or

Sequoia and Kings Canyon National Parks (Figures 11, 14, and 16). The ordinations

always position my flora on the east edge of the Westside group, suggesting that my flora is

closely related to the eastside floras with some westerly characteristics (Figures 12, 15, and

17).

83 84

By overlaying a set of environmental variables onto the ordinations, I found that the floras' locations, geographical distance from the San Joaquin Roadless Area, and their numbers of taxa are the most important factors in explaining variation among floras. My results support those of Taylor(1977) where 85% of the variation among floras was explained by geographical distance; however, these are not the only factors contributing to these differences. Elevational range and aerial extent are also weakly correlated with floristic composition. In addition, I agree with Bowers and McLaughlin (1982) who found that collecting time and the presence of aquatic habitats and canyon environments affect relative species richness and can explain some of the variation among floras. For example, many Northern Hemisphere taxa are widespread aquatic plants. Lassen National Park and the San Bernardino Mountains, both of which have several lakes, have the largest proportions of these taxa. My study area, which contains no lakes, has a much smaller proportion of Northern Hemisphere taxa.

The geographically close Sierran floras are shown to be similar in composition and affinities. Floras farther away from the San Joaquin Roadless Area show the long distance relationships of the Sierran floras. In general, however, the floras are similar by geographic zones. The Sweetwater Mountains, Upper Walker River Watershed, Bodie Hills, and

White Mountains were consistently grouped together as Great Basin floras, and all show the greatest proportion of Great Basin taxa (Figures 11, 14, and 16). I had expected the

San Bernardino Mountains to be grouped with these because of the large number of desert taxa coded as Great Basin. However, the predominance of Lowland California taxa in the middle and lower elevations and the, albeit depauperate, Sierran flora at upper elevations

(Krantz 1994, Taylor 1977) grouped this flora with the Sierran and northern floras instead of the Great Basin (Figure 11). 85

The Trinity Alps were consistently classified as a northern and western flora in the analyses, due to a high percentage of Circumboreal taxa, despite low elevations and limited high elevation habitats, together with a high percentage of Lowland California taxa.

Analyses of Phytogeographic Affinities

My analyses agreed with those of Stebbins' (1982). He showed a relationship between the phytogeographic affinities of Sierran taxa and their distribution according to moisture availability. Given that most new migrants into the study area came from the closely surrounding Sierran flora, the availability of favorable conditions influenced the relative success of colonizing taxa with different origins and adaptations (Bliss 1985, Went

1948). The proportions of taxa from each region of origin are reflected in the dominating

moisture regimes of the area (Stebbins 1982). The pattern is enhanced by recognizing the

dominating environment of each elevational zone as suggested by the climate data (Table

1).

My area has a much larger proportion of Old Cordilleran taxa than does the entire

Sierra Nevada (Table 11). These taxa are adapted to relatively dry environments compared

to Circumboreal taxa, but to wetter environments than Great Basin or Lowland California

taxa (Stebbins 1982). My upper montane and subalpine zones, dominated by forests and

subalpine meadows, make up the majority of the area. Here a deep snowpack and only

moderately deep pumice create conditions of moderate water stress, allowing a larger

proportion of Old Cordilleran taxa to find acceptable habitat within the study area as

compared to the Sierra as a whole.

Circumboreal taxa make up a substantially lower proportion of my flora compared

to that of the entire Sierra. These taxa are adapted to the most mesic habitats, such as wet

alpine meadows, streamsides, and lakeshores (Stebbins 1982, 1997 Pers. comm.). These 86 are very limited within my area. In addition, the proportion of these taxa does not increase in my alpine as in the entire Sierran alpine (Table 12). The relatively low elevation of my alpine (3536 m) and the dominance of windy, dry, exposed ridges have limited available habitat for species diversity in this affinity category.

The proportion of Great Basin taxa in my area is lower than that of the entire Sierra

(Table 11). I did not expect this result because dry, exposed alpine and large expanses of open pumice where water stress is very high seemed to characterize the study area.

However, areas dominated by drought-adapted Great Basin taxa, are mostly located close to the volcanic vents in the montane zone. My subalpine zone, the majority of the study area, contains a very low proportion of Great Basin plants (Table 13). In the alpine, the proportion of Great Basin taxa increases, but not enough to offset the moderating effect of the subalpine environment (Table 13). As a result, species diversity in this affinity category is reduced compared to that of the entire Sierra (Table 11).

There is a smaller proportion of Lowland California taxa in my area compared to the entire Sierra due to the lack of extensive montane area and its eastern location (Table 11).

These taxa are usually found in dry sites at lower elevations, and as expected, their proportion of the flora decreases as elevation increases (Stebbins 1982). However, in my alpine the proportion of these taxa is double that of the entire Sierran alpine (Table 12). The overall lower elevation and predominance of exposed, scree covered ridges in my alpine create the warm, xeric habitats suitable for Lowland California taxa. These habitats are generally limited in the Sierran alpine where Lowland California taxa are mostly limited to areas disturbed by needle ice (Bliss 1985). 87

Mammoth Gap as a Migration Corridor

My results confirm the Mammoth Gap as an effective migration corridor for

Lowland California taxa to the east side of the Sierra. When compared to other eastside floras, my area has the highest proportion of Lowland California taxa, besides Mammoth

Mountain (Figure 20). These taxa originated in cis-montane California and have migrated upward and eastward with the rise of the Sierra (Stebbins 1982). They are generally found at middle to low elevations on the west side and in warm spots in the higher elevations, but several have crossed the range and found suitable habitat on the east side. The higher proportion on Mammoth Mountain compared to the study area suggests that Mammoth

Pass, the lowest elevation gap, is a more effective corridor than the higher elevation

Deadman Pass.

Origins of the San Joaquin Roadless Area Flora

The recovery of the area's flora after the volcanic disturbance has occurred rather quickly. High winds over the crest and many seed sources from areas outside the blast zone contributed to this rapid recovery. Heavy snowfall and fast-draining soils created the perfect environment for the red fir forest from the west side of the Sierra to colonize large parts of the area (Oosting and Billings 1943). Wet areas provided habitat for taxa from the surrounding high Sierra to take hold. Glass Creek Meadow shows surprisingly high productivity despite disturbance (Millar and Woolfenden in press). Dry pumice areas provided habitat for Great Basin and drought adapted Sierran taxa.

Migration into the study area was not limited to the close surrounding locations, as documented by the several taxa with range extensions in the area. Since the disturbance, taxa from as far away as the Donner Pass area; the White, lnyo, and other desert ranges; as well as the southern and western Sierra have arrived and found suitable habitat here. 88

Immigration continues today, as evidenced by several taxa that occur as single, small populations in the area. Those occurring in dry, exposed sites, such as Descuriana incana, Cirsium scariosum, and Tetradymia canescens, may represent the most recent immigrants. The single tree of Populus balsamifera ssp. trichocarpa is also a recent immigrant, but probably human assisted. Other taxa with limited distribution in the area are found in high elevation wet sites or on the limestone, but they may be restricted by limited habitat.

The Origin of Arabis pinzlae

One taxon did not migrate here but rather originated here, possibly very recently.

Despite the recent volcanic disturbance of the area, evidence points to a Sierran origin for the rare Arabis pinzlae. Its seeds are winged for effective wind dispersal. Prevailing winds in the area blow from west to east, creating a perfect vector for seed dispersal to the White

Mountains. Boundary Peak, the location of the disjunct White Mountain populations, lies directly east of the ridge where the Sierran A. pinzlae grows.

Seed dispersal between the White Mountains and the Sierra is highly probable due to the short distance between the two ranges (Stebbins 1982). The major direction of plant exchange between the Sierra Nevada and the White Mountains is usually from the east to the west, due to the lack of suitable habitat in the White Mountains for Sierran plants

(Reveal 1979). However in this case, the high-elevation shifting scree environment, to which A. pinzlae is adapted (Morefield 1994), exists in both ranges. Winged seeds; fully dehiscent, upright fruits; late season wind direction; and the physical locations of the disjunct populations support a west to east migration scenario. The taxon's restricted distribution in the Sierra and scattered distribution in the White Mountains also supports my hypothesis of wind dispersal to the White Mountains. 89

Interestingly, unless this Sierran ridgetop was protected from the recent volcanics, this taxon had only about 600 years to evolve and disperse. Although the ridge where it grows is not covered in a blanket of pumice, there are pumice grains in the soil indicating that tephra fall did occur here, probably killing the plant life (Millar 1999 pens. comm.).

The pumice is very quickly eroded by wind and water, especially from exposed ridgetops, therefore little evidence of disturbance remains in these areas.

However, there is a possibility that parts of the high alpine ridges may not have been directly affected by the volcanic blasts. Dendrochronological evidence shows tree death on San Joaquin Ridge at the same time as the eruptions, but confounding factors of climate change and incomplete mapping and understanding of the volcanic blasts and tephra flow leave open the possibility of plant survival in the high alpine (Millar 1999 Pers. comm.).

Considering this possibility in light of recent vegetation studies on 14-year old devastated areas of Mount St. Helens (Titus etal. 1998), wind dispersed plants from possible refugia in the high alpine of San Joaquin Ridge may have been some of the first taxa to colonize the lower elevations of the study area. Some 50 taxa (11 % of the flora) occur in the alpine zone as well as in the subalpine and montane zones (Appendix A). Many of these taxa are of Great Basin and Old Cordilleran affinity and are well adapted to the dry, pumice habitats available after the eruptions. Twelve of these are listed among the early colonizers of the primary successional habitats on Mount St. Helens (Table 15). One large- seeded, non-wind dispersed taxon, Lupinus lepidus, is believed to have survived in a few small patches within the blast zone of the Mount St. Helens eruption (Bishop 1996). The survival of this plant on Mount St. Helens and our inadequate understanding of the destructiveness of the recent Ingo eruptions strongly suggest the possibility of survivors in high alpine refugia in the study area. Prevailing west winds and the tenacity of L. lepidus, 90 of which my flora contains three varieties, point to the taxa listed in Table 15 as some of the first colonizers of the blast zone in the San Joaquin Roadless Area. 91

Table 15. Taxa that occur on the Pumice Plain on Mount St. Helens (Titus et al. 1998) and also in all three elevation zones of the San Joaquin Roadless Area. If plants in the high alpine were not destroyed during the recent Ingo Craters eruptions, they may be some of the first colonizers of the study area.

Achnatherum occidentale ssp. occidentale Carex spectabilis Agoseris glauca Elymus elymoides ssp. elymoides Antennaria rosea Eriophyllum lanatum Arnica nevadensis Juncus parryi Calyptridium umbellatum Lupinus lepidus Carex leporinella Trisetum spicatum CONCLUSIONS

The cluster analyses, ordinations, and affinity analyses for the 13 floras and subsets show that the San Joaquin Roadless Area is floristically centered in relationship to other

California mountain floras. As predicted, the Trinity Alps and San Bernardino Mountains are distant floristically as they are geographically. Overall the floras most similar to the San

Joaquin Roadless Area are geographically close (Tuolumne Meadows, Hall Research

Natural Area, and Mammoth Mountain).

Floristically the area is more similar to the west side of the Sierra Nevada, rather than to eastside floras. However it is closer to eastside floras than any Westside flora is, so it is floristically centered in the east/west direction, as well as from north to south. In certain analyses it is grouped with the eastern group. This position for the San Joaquin

Roadless Area may be due to the stronger Great Basin relationships shown by the affinities of the montane and alpine zones, whereas the affinities of the subalpine zone, which is the main core of the area, maintain a strong relationship to the west side of the Sierra Nevada.

I would stress the importance of preserving this relatively pristine example of the flora of the central eastern Sierra. Its flora has had a short time to develop, and in this time two rare plants and several plants exhibiting range extensions have established here. The area contains a large and unique eastside red fir forest and is habitat to fur-bearing mammals and rare birds and amphibians.

The protection of the two rare and endangered plant taxa here deserves special consideration. The impacts of off-trail mountain bikes and hikers on the White Wing population of Lupinus duranii need to be monitored and reduced. The Sierran population of

Arabis pinzlae does not receive the wilderness protection of those in the White Mountains.

Efforts must be made to protect its habitat from human impacts in order to preserve this rare and newly developed element of the genetic diversity of Arabis.

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Titus, J.H., S. Moore, M. Arnot, and P.J Titus. 1998. Inventory of the vascular flora of the blast zone, Mount St. Helens, Washington. Madroño 45:146-161.

USDA Forest Service. 1988. Land and Resource Management Plan, Ingo National Forest. Pacific Southwest Region, USDA Forest Service, San Francisco, CA.

USDA Forest Service. 1995. Existing and Historic Conditions of the Mammoth to June Area: Preliminary Results of the Mammoth to June Ecosysten Analysis. Ingo National Forest, Bishop, CA.

USDA Forest Service. 1997. Desired Condition for the Mammoth to June Area: The second of three reports for the Mammoth to June Ecosystem Analysis. Ingo National Forest, Bishop, CA.

USDI National Park Service. 1996. A plant checklist of Devils Postpile National Monument, Madera County, California. Unpublished checklist compiled by the Devils Postpile National Monument, Madera County, CA.

Weeden, N.F. 1986. A Sierra Nevada Flora. Wilderness Press, Berkely, CA.

Went, F.W. 1948a. Some parallels between desert and alpine flora in California. Madroño 9:241-249.

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Woolfenden, W., T. Higley, and T. Hicks. 1995. Vegetation. Pages V-1 to V-48 in USDA Forest Service, author. Existing and Historic Conditions of the Mammoth to June Area: Preliminary Results of the Mammoth to June Ecosysten Analysis. Ingo National Forest, Bishop, CA.

Woolfenden, W. 1996. Quaternary vegetation history. Pages 47-70 in Status of the Sierra Nevada, Volume II: Assessments and scientific basis for management options. Sierra Nevada Ecosystem Project, Final report to Congress. Wildland Resources Center Report No. 37, University of California, Davis, CA.

Wulff, E.V. 1950. An introduction to historial plant geography. Chronica Botanica Company, Waltham, MA. APPENDIX A. Annotated species list of the San Joaquin Roadless Area. Collection numbers are in parentheses after validating authors name. Phytogeographic affinity of each taxon (Stebbins 1997) is noted at the end of each entry. Specimens are deposited at HSC.

FERNS AND FERN ALLIES

DENNSTAEDTIACEAE

Pteridium aquilinum (L.) Kuhn var. pubescens L. Underw. (142, 760) Occasional in open forests; 2400 - 3000 m. Northern Hemisphere.

DRYOPTERIDACEΑE

Cystopteris fragilis (L.) Bernh. (330, 468, 602, 832, 894, 903, 1012) Common in rocks; 2400 - 3400 m. Northern Hemisphere.

EQUISETACEAE

Equisetum arvense L. (24, 887) Common along stream sides; 2400 - 3000 m. Northern Hemisphere.

OPHIOGLOSSΑCEΑE

Botrychium simplex Hitchc. (1234) One small population near the middle of Glass Creek Meadow; 2530 m. Northern Hemisphere.

PTERIDACEAE

Cryptograma acrostichoides R. Br. (968) Occasional in shaded rocks; 2600 - 3000 m. Old Cordilleran.

Pellaea breweri D. Eaton (697, 904, 1216) Occasional in rocks; 2400 - 3400 m. Old Cordilleran.

SELΑGINELLΑCEΑE

Selaginella watsonii L. Underw. (1060, 1192) Common on the limestone and near the summit of San Joaquin Mountain; 3000 - 3536 m. Lowland California.

GYMNOSPERMS

PINACEAE

Abies magnifica Andr. Murray var. magnifica (159) Abundant in montane and subalpine forests; 2400 - 3000 m. Old Cordilleran.

Pinus albicaulis Engelm. (191, 871) Common in montane and subalpine forests and on high ridges; 2400 - 3400 m. Old Cordilleran.

106 107

Pinus contorta Loudon ssp. murrayana (Grey. & Balf.) Critchf. (158) Abundant in montane and subalpine forests; 2400 - 3000 m. Old Cordilleran.

Pinus flexilis James (983) One population near the summit of North Point; 2970- 2987 m. Old Cordilleran.

Pinus jeffreyi Grey. & Balf. (157) Abundant in montane forests and two trees elevationally disjunct on the limestone outcrop; 2400 - 3000 m. Old Cordilleran.

Pinus monticola Douglas (156) Common in montane and subalpine forests; 2400 - 3000 m. Old Cordilleran.

Tsuga mertensiana (Bong.) Carriēre (296, 433) Occasional in subalpine forests and in protected areas at higher elevations; 2600 - 3400 m. Old Cordilleran.

DICOTS

APIACEAE

Angelica lineariloba A. Gray (153) Common in montane chaparral and dry forests and elevationally disjunct on the limestone outcrop; 2400 - 3000 m. Lowland California.

Cicuta maculata L. var. angustifolia Hook. (990) 2400 - 2600 m. Occasional along streams in the red fir forest. Lowland California.

Cymopteris terebinthinus (Hook.) Μ.E. Jones var. petraeus (Μ.E. Jones) Goodrich (239, 432, 518, 664) Common on open pumice slopes, exposed ridges, and in montane chaparral. 2600 - 3000 m. Great Basin.

Heracleum lanatum Michaux (223, 234, 640) Occasional in dry edges of meadows, open aspen stands and riparian areas; 2600 - 3000 m. Northern Hemisphere.

Liguisticum grayii J. Coulter & Rose (784, 833, 991) Common in open forests and avalanche areas; 2400 - 3400 m. Circumboreal.

Osmorhiza chilensis Hook. & Am. (1011) Common in open subalpine forests and scrubs; 2600 - 3000 m. Lowland California.

Osmorhiza occidentalis (Nutt.) Torrey (283) Common in open subalpine forests and scrubs; 2600 - 3000 m. Old Cordilleran.

Perideridia parishii (J. Coulter & Rose) Nelson & J.F. Macbr. ssp. latifolia (A. Gray) Chuang & Constance (20, 69, 117, 231, 746, 1207) Abundant in wet meadows and riparian areas; 2400 - 3400 m. Lowland California.

Sphenosciadium capitellatum A. Gray (22, 260) Occasional in riparian areas and on exposed ridges especially near Deadman Pass; 2400 - 3400 m. Old Cordilleran. 108

APOCYNACEAE

Apocynum androsaemifolium L. (150, 173, 669) Common understory plant in mixed forests and open sagebrush in the avalanche zone; 2400 - 3000 m. Northern Hemisphere.

ASTERACEAE

Achillea millefolium L. var. lanulosa (Nutt.) Piper (57) Common in wet meadows, especially Glass Creek Meadow; 2600 - 3000 m. Old Cordilleran.

Ageratina occidentalis (Hook.) R. King & H. Robinson (703) Occasional on rock outcrops; 2600 - 3000 m. Old Cordilleran.

Agoseris aurantiaca (Hook.) E. Greene (635) Occasional in open sagebrush; 2600 - 3000 m. Old Cordilleran.

Agoseris elata (Nutt.) E. Greene (1218) Occasional in montane chaparrals; 2600 - 3400 m. Lowland California.

Agoseris glauca (Pursh) Raf. var. laciniata (D. Eaton) F.J. Smiley (444) Occasional in open pumice, especially at Crater Flat; 2400 - 2600 m. Old Cordilleran.

Agoseris glauca (Pursh) Raf. var. monticola (E. Greene) Q. Jones (48, 92, 202, 322, 463) Common in open pumice; 2400 - 3400 m. Old Cordilleran.

Agoseris retrorsa (Benth.) E. Greene (626, 875) Occasional in open scrubs at the edge of the red fir forest; 2600 - 3000 m. Lowland California.

Antennaria corymbosa E. Nelson (83, 97, 1131) Common in dry meadow edges; 2600 - 3400 m. Old Cordilleran.

Antennaria dimorpha (Nutt.) Torrey & A. Gray (443) Occasional in pumice at Crater Flat; 2480 m. Great Basin.

Antennaria media E. Greene (1041) Occasional in the alpine fell fields near the summit of San Joaquin Peak; 3400 - 3536 m. Old Cordilleran.

Antennaria rosea E. Greene ssp. confinis (E. Greene) R. Bayer (360, 440) Common on exposed ridges and open pumice areas; 2400 - 3400 m. Old Cordilleran.

Antennaria rosea E. Greene ssp. rosea (218, 692, 902) Common in forest openings; 2400 - 3000 m. Old Cordilleran.

Arnica amplexicaulis Nutt. (1115, 1179) Occasional in periodically dry riparian areas; 2600 - 3400 m. Old Cordilleran.

Arnica chamissonis Less. ssp. foliosa (Nutt.) Maguire (233, 528, 908) Common in wet meadows; 2600 - 3000 m. Old Cordilleran. 109

Arnica latifolia Bong. (845) Occasional along streams; 2600 - 3000 m. Old Cordilleran.

Arnica mollis Hook. (303, 835) Occasional along streams; 2600 - 3000 m. Old Cordilleran.

Arnica nevadensis A. Gray (598, 693, 709, 955, 1006) Common in subalpine forests, on open slopes and ridges, and in lower elevation forest openings; 2400 - 3400 m. Old Cordilleran.

Arnica parryi A. Gray (1144, 1169) Occasional along streams; 3000 - 3400 m. Old Cordilleran.

Artemisia arbuscula Nutt. ssp. arbuscula (312, 351, 572, 716, 763) Abundant on exposed ridges and south facing steep slopes; 2600 - 3400 m. Great Basin.

Artemisia arbuscula Nutt. ssp. thermopola Beetle (853) Occasional in open mixed subalpine forest; 2600 - 3000 m. Great Basin.

Artemisia douglasiana Sesser (257, 680) Common along streams and among open sagebrush in avalanche zone; 2400 - 3000 m. Great Basin.

Artemisia ludoviciana Nutt. ssp. candicans (Rydb.) Keck (1224) Range extension to the south, first report for Mono County. Occasional along stream running through and below Minaret Meadow; 2690 m. Great Basin.

Artemisia ludoviciana Nutt. ssp. incompacta (Nutt.) Keck (966) Occasional along streams in the red fir forest; 2400 - 2600 m. Great Basin.

Artemisia ludoviciana Nutt. ssp. ludoviciana (965, 1183) Occasional along streams in the red fir forest and also on the limestone; 2400 - 3000 m. Great Basin.

Artemisia michauxiana Sesser (987, 1180) First documented report for the Sierra Nevada. Occasional along streams in the red fir forest and in the avalanche zone; 2400 - 3000 m. Great Basin.

Artemisia tridentata Nutt. ssp. vaseyana (Rydb.) Beetle (47, 252, 629, 668, 670, 681, 690, 720, 776, 974, 1071, 1080, 1081, 1086) Abundant in the montane chaparrals, mixed forest understory, and avalanche areas and on mid-elevation exposed ridges; 2400 - 3000 m. Great Basin.

Aster alpigenus (Torrey & A. Gray) A. Gray var. andersonii (A. Gray) M. Peck (70, 90, 529, 1111) Abundant in wet meadows; 2600 - 3000 m. Old Cordilleran.

Aster breweri (A. Gray) Semple (596, 683, 694, 1007) Occasional on open slopes and in forest openings; 2400 - 3400 m. Lowland California.

Aster integrifolius Nutt. (291, 731, 1027) Occasional in dry meadows, especially upper Glass Creek Meadow; 2600 - 3000 m. Old Cordilleran. 110

Aster occidentalis (Nutt.) Torrey & A. Gray var. occidentalis (851, 1104) Occasional in dry meadows; 2600 - 3000 m. Old Cordilleran.

Aster scopulorum A. Gray (975) Occasional in rocks; 2600 - 2800 m. Great Basin.

Chaenactis alpigena Sharsm. (266, 1015) Occasional in open pumice areas; 2600 - 3000 m. Old Cordilleran.

Chaenactis douglasii (Hook.) Hook. & Am. var. alpina A. Gray (1057) Rare in California, CLAPS List 2. Occasional in the alpine fell fields near the summit of San Joaquin Peak; 3400 - 3536 m. Great Basin.

Chaenactis douglasii (Hook.) Hook. & Arn. var. douglasii (54, 175, 989, 1036) Common in open pumice and understory of mixed Jeffrey pine forest; 2400 - 3000 m. Great Basin.

Chrysothamnus nauseosus (Pallas) Britton ssp. albicaulis (Nutt.) H.M. Hall & Clements (310, 765) Common among rocks in montane chaparral; 2600 - 3000 m. Great Basin.

Chrysothamnus parryi (A. Gray) E. Greene ssp. asper (E. Greene) H.M. Hall & Clements (41, 169, 1067, 1075, 1077, 1078, 1084) Abundant in open pumice areas and in the understory of open mixed Jeffrey pine forests; 2400 - 3000 m. Great Basin.

Chrysothamnus parryi (A. Gray) E. Greene ssp. monocephalus (Nelson & Kenn) H.M. Hall & Clements (574) Common on exposed ridges; 3000 - 3500 m. Great Basin.

Chrysothamnus viscidflorus (Hook.) Nutt. ssp. puberulus (D.C. Eaton) H.M. Hall & Clements (208, 785, 1118) Common on exposed ridges; 2600 - 3500 m. Great Basin.

Cirsium scariosum Nutt. (1114) One small population in an intermittent streambed that feeds into the north canyon of Deadman Creek; 2840 m. Great Basin.

Crepis acuminate Nutt. (319, 568, 898, 1120) Common on exposed ridges; 2600 - 3400 m. Great Basin.

Ericaineria bloomeri (A. Gray) J.F. Macbr. (168, 854) Occasional in dry forest edges and openings; 2400 - 3000 m. Great Basin.

Ericameria discoidea (Nutt.) G. Nesom (264) Common in open pumice, dry meadow edges, and on exposed ridges; 2600 - 3400 m. Great Basin.

Ericameria nana Nutt. (377, 378) Occasional in dry forest edges and openings; 2600 - 3000 m. Great Basin. 111

Ericameria suffruticxsa (Nutt.) G. Nesom (567) Occasional on exposed ridges; 3000 - 3536 m. Great Basin.

Erigeron algidus Jepson (808, 957) Occasional on exposed ridges and in high elevation drainages; 3000 - 3536 m. Old Cordilleran.

Erigeron barbellulatus E. Greene (50, 201, 339, 457, 787, 1038, 1039, 1056, 1189) Common in open pumice and on exposed ridges and peaks; 2400 - 3536 m. Old Cordilleran.

Erigeron breweri A. Gray var. breweri (174, 809) Occasional in open rocky areas and in dry forests; 2400 - 3400 m. Old Cordilleran.

Erigeron compositus Pursh. (713, 792) Common on exposed ridges and alpine fell fields; 3000 - 3536 m. Old Cordilleran.

Erigeron coulteri Porter (247, 623, 1000) Common in wet meadows; 2400 - 3000 m. Old Cordilleran.

Erigeron linearis (Hook.) Piper (459) Occasional in pumice at Crater Flat; 2400 - 2600 m. Old Cordilleran.

Erigeron peregrinus (Pursh) E. Greene var. callianthemus (E. Green) Cronq. (68, 119, 241, 243, 285, 493, 556, 834, 954, 1088) Common in meadows; 2400 - 3000 m. Old Cordilleran.

Erigeron peregrinus (Pursh) E. Greene var. hirsutus Cronq. (822) Occasional in riparian areas; 2600 - 3000 m. Old Cordilleran.

Eriophyllum lanatum (Pursh) James Forbes var. integrifolium (Hook.) F.J. Smiley (38, 129, 212, 323, 452, 571, 988) Common in open pumice and on exposed ridges; 2400 - 3400 m. Lowland California.

Hazardia whitneyi (A. Gray) E. Greene var. whitneyi (707) Occasional on exposed ridges; 3000 - 3400 m. Old Cordilleran.

Hieracium albiflorum Hook. (222, 685) Occasional in the red fir and mixed Jeffrey pine forest; 2600 - 3000 m. Old Cordilleran.

Hieracium gracile Hook. (1087) Occasional in red fir forest; 2600 - 2700 m. Old Cordilleran.

Hieracium horridum Fries (187, 304) Occasional on pumice slopes; 2600 - 3400 m. Old Cordilleran.

Hulsea vestita A. Gray ssp. vestita (55, 198, 1079) Common in open pumice and on exposed ridges; 2600 - 3500 m. Lowland California.

Machaeranthera canescens (Pursh) A. Gray) var. canescens (294, 578, 900) Occasional in dry meadows and on exposed ridges; 2600 - 3400 m. Great Basin. 112

Raillardella argentea (A. Gray) Α. Gray (51, 358, 429, 542) Occasional in open pumice and on exposed ridges; 2600 - 3536 m. Old Cordilleran.

Raillardella scaposa (Α. Gray) Α. Gray (570) Occasional on high elevation open slopes and exposed ridges; 3000 - 3400 m. Lowland California.

Senecio canus Hook. (397, 428, 762, 810, 1063) Common in open pumice, rocky and scree areas; 3000 - 3400 m. Old Cordilleran.

Senecio integerrimus Nutt. var. exaltatus (Nutt.) Cronq. (689) Occasional in forest openings; 2600 - 3000 m. Old Cordilleran.

Senecio scorzonella E. Greene (102, 240, 520, 1148) Common in wet to dry meadows; 2600 - 3400 m. Old Cordilleran.

Senecio spartoides Torrey & Α. Gray (143, 255) Occasional in mixed Jeffrey pine forest; 2400 - 3000 m. Great Basin.

Senecio triangularis Hook. (2, 165, 244, 1101, 1203) Common in wet meadows and along streams; 2400 - 3000 m. Old Cordilleran.

Senecio werneriifolius A. Gray (793, 1054, 1055) Common in alpine fell fields and on exposed ridges; 3400 - 3536 m. Old Cordilleran.

Solidago canadensis L. ssp. elongata (Nutt.) Keck (374, 823) Occasional in riparian areas; 2600 - 3000 m. Northern Hemisphere.

Stephanomeria spinosa (Nutt.) Tomb (1018) Occasional. Found only on the bench between the Glass and Deadman watersheds; 2800-2840 m. Old Cordilleran.

Stephanomeria tenuifolia (Torrey) H. M. Hall (132, 177, 215, 1230) Common on pumice soils in montane forests and elevationally disjunct on the limestone outcrop; 2400 - 3000 m. Old Cordilleran.

Taraxacum officiηale Wigg. (101, 506, 658) Occasional in Glass Creek Meadow; 2600 - 3000 m. Alien, introduced from Europe.

Tetradymia canescens DC. (913) One population in the quartz latite rocks on the summit of June Mountain; 3083m. Great Basin.

Wyethia mollis A. Gray (320, 449) Occasional on open slopes and exposed ridges; 2600 - 3000 m. Lowland California.

ΒEΤULΑCEΑE

Alnus incana (L.) Moench ssp. tenuifolia (Nutt.) Breitung (769) Common only in stream courses in the south canyon of Deadman Creek; 2600 - 3000 m. Old Cordilleran. 113

BORAGINACEAE

Cryptantha ambigua (A. Gray) E. Greene (178, 220, 484, 645) Common in mixed Jeffrey pine forest and open sagebrush; 2400 - 3000 m. Great Basin.

Cryptantha circumscissa (Hook. & Arn.) I.M. Johnston (1059) Occasional in alpine fell fields; 3400 - 3536 m. Great Basin.

Cryptantha echinella E. Greene (1221) Occasional, found only in open forests on North Point; 2600 - 3000 m. Great Basin.

Cryptantha glomeriflora E. Greene (1164) Occasional; found only in bowl below San Joaquin Peak; 3200 m. Great Basin.

Cryptantha humilis (E. Greene) Payson (476) Occasional on pumice in the Jeffrey pine forest; 2400 - 2600 m. Great Basin.

Cryptantha nubigena (E. Greene) Payson (196, 361, 402, 1083) Common in pumice and on exposed ridges; 2600 - 3400 m. Great Basin.

Cryptantha sobolifera Payson (1037) Occasional in the rocks on San Joaquin Peak; 3500 - 3536 m. Great Basin.

Hackelia micrantha (Eastw.) J. Gentry (7, 137, 162, 641, 927, 976) Common in dry meadow edges and montane chaparral; 2400 - 3000 m. Old Cordilleran.

Plagiobothrys cognatus (E. Greene) I.M. Johnston (515) Occasional in Glass Creek Meadow; 2400 - 3000 m. Old Cordilleran.

Plagiobothrys hispidulus (E. Greene) I.M. Johnston (543, 918) Common in Glass Creek Meadow; 2600 - 3000 m. Lowland California.

Plagiobothrvs hispidus A. Gray (388, 399, 882) Abundant in mixed Jeffrey pine forest and dry meadow areas; 2600 - 3000 m. Lowland California.

BRASSICACEAE

Arabis xdivericarpa Nelson (967) Occasional along streams in red fir forest; 2400 - 2600 m. Great Basin.

Arabis drummondii A. Gray (225, 888, 952, 999, 1113) Common in dry meadow and forest edges; 2400 - 3000 m. Old Cordilleran.

Arabis holboellii Horner. var. pinetorum (Tidestrom) Rollins (394, 396, 424, 435, 610, 1108) Common in mixed forests and open pumice; 2400 - 3000 m. Old Cordilleran.

Arabis holboellii Horner. var. retrofracta (Graham) Rydb. (441, 1003) Occasional in open pumice and red fir forest; 2400 - 2600 m. Old Cordilleran. 114

Arabis lemmonii S. Watson var. depauperate (Nelson & Kenn.) Rollins (481, 1105, 1152, 1227) Occasional in open pumice and on the limestone; 2400 - 3040 m. Old Cordilleran.

Arabis lemmonii S. Watson var. lemmonii (136, 370, 478, 1185) Common in dry forests, open pumice and on exposed ridges; 2600 - 3400 m. Old Cordilleran.

Arabis lyalii S. Watson var. nubigena (J.F. Macbr. & Payson) Rollins (367) Occasional on exposed ridges and open slopes; 3000 - 3400 m. Old Cordilleran.

Arabis pinzlae Rollins (1158) RARE CLAPS List 1B, first record for the Sierra Nevada. Widely scattered among whitebark pine krummholz on southeast-facing side of the east trending spur ridge below Two-Teats. 3200 - 3280 m. Great Basin.

Arabis platysperma A. Gray var. howellii (S. Watson) Jepson (130, 483, 803) Occasional on exposed ridges, open slopes and pumice; 2400 - 3400 m. Old Cordilleran.

Arabis platysperma A. Gray var. platysperma (149, 213, 292, 307, 420, 431, 589, 712, 1194, 1231) Common in dry areas throughout; 2400 - 3400 m. Old Cordilleran.

Arabis puberula Torrey & A. Gray (876, 1066) Occasional on open metamorphic rock slopes and scree; 3000 - 3536 m. Great Basin.

Arabis repanda S. Watson ssp. repanda (1008) Occasional in red fir forest; 2400 - 2600 m. Great Basin.

Barbarea orthoceras Ledeb. (500, 501) Occasional along streams; 2400 - 2600 m. Northern Hemisphere.

Cardamine breweri S. Watson var. breweri (5, 1030) Common in wet meadows; 2600 - 3000 m. Old Cordilleran.

Descuriana calίfornίca (A. Gray) O. Schulz (151, 164, 581, 905, 1016) Occasional in mixed forests and on open slopes; 2400 - 3400 m. Northern Hemisphere.

Descuriana incana (Fischer & C. Meyer) Dorn (959) One small population on an exposed basalt outcrop on San Joaquin Ridge at the southeast edge of Deadman Pass; 2980 m. Northern Hemisphere.

Descuriana incisa (A. Gray) Britton var. incise (490, 1110) Occasional along streams; 2400 - 3000 m. Old Cordilleran.

Draba albertina E. Greene (408, 735, 742, 881, 947, 1138, 1160, 1213) Common in wet meadows and in riparian areas; 2600 - 3400 m. Old Cordilleran.

Draba breweri S. Watson (336) Occasional in rock crevices on exposed ridges; 3000 - 3536 m. Old Cordilleran. 115

Erysimum capitatum (Douglas) E. Greene ssp. perenne (Coy.) R.A. Price (133, 154, 251, 300, 391, 628) Common in dry forests; 2400 - 3000 m. Lowland California.

Lepidium virginicum L. var. virginicum (111 2) One small population in the now closed upper Deadman Road, just across the creek; 2525 m. Northern Hemisphere, although probably alien in California (Hickman 1993).

Phoenicaulis cheiranthoides Torrey & A. Gray (293, 711) Occasional on exposed ridges; 2600 - 3400 m. Old Cordilleran.

Rorippa curvipes E. Greene var. curvipes (1222) Occasional in wet meadows; 2600 - 3000 m. Northern Hemisphere.

Rorippa curvisiliqua (Hook.) Britton (970) Occasional in aspen near standing water; 2400 - 2600 m. Circumboreal.

Rorippa nasturtium-aquaticum (L.) Hayek (289) Common in wet meadows, widespread in Glass Creek Meadow; 2600 - 3000 m. Alien, introduced from Europe.

Rorippa sinuata (Torrey & A. Gray) A. Hitchc. (1097) Occasional in wet meadows, found only in Minaret Meadow; 2600 - 3000 m. Northern Hemisphere.

Streptanthus tortuosus Kellogg var. orbiculatus (E. Greene) H.M. Hall (356, 590, 777) Common on open slopes; 2600 - 3400 m. Lowland California.

CAPRIFOLIACEAE

Lonicera involucrata (Richardson) Banks var. involucrata (981, 1103) Occasional along streams in the red fir and lodgepole pine forests; 2600 - 3000 m. Old Cordilleran.

Sambucus racemosa L. var. microbotrys (Rydb.) Kearney & Peebles (1, 249, 637) Common in open riparian areas; 2600 - 3000 m. Old Cordilleran.

Symphoricarpos rotundifolius A. Gray var. rotundifolius (145, 261, 548, 580, 625, 698, 915, 950, 1065) Abundant in montane chaparrals, aspen thickets, and forests; 2400 - 3400 m. Old Cordilleran.

CARYOPHYLLACEAE

Arenaria kingii (S. Watson) Μ.E. Jones var. glabrescens (S. Watson) Maguire (43, 328, 1043, 1201) Common in open pumice and on exposed ridges and open slopes; 2400 - 3400 m. Old Cordilleran.

Minuartia nuttallii (Pax) Briq. ssp. gracilis (Robinson) McNeill (184, 197, 417, 801, 1175) Common on open pumice, bowls and slopes; 2600 - 3536 m. Old Cordilleran. 116

Sagina saginoides (L.) Karsten (733, 1028, 1093) Common in wet meadows; 2600 - 3000 m. Northern Hemisphere.

Silene bernardina S. Watson (771, 1184) Occasional on rocky slopes and on the limestone; 2600 - 3000 m. Lowland California.

Silene sargentii S. Watson (203, 369, 802, 963) Occasional on open slopes and exposed ridges; 3000 - 3536 m. Old Cordilleran.

Stellaria calycantha (Ledeb.) Bong. (1021) Occasional in wet meadows; 2600 - 3000 m. Old Cordilleran.

Stellaria crispa Cham. & Schldl. (837, 1162, 1193) Occasional in riparian areas; 3000 - 3300 m. Old Cordilleran.

Stellaria longipes Goldie var. longipes (76, 84, 49'7, 536, 895, 907) Abundant in wet meadows; 2400 - 3000 m. Northern Hemisphere.

Stellaria umbellate Karelin & Kir. (1033, 1134, 1214) Occasional in wet meadows and riparian areas; 2600 - 3536 m. Old Cordilleran.

CHENOPODIACEAE

Chenopodium atrovirens Rydb. (284, 545, 1119) Weedy and dominant in sheep bedding areas on the north side of lower Glass Creek Meadow and on the bench between Glass and Deadman watersheds; 2600 - 3000 m. Old Cordilleran.

ERICACEAE

Arctostaphylos nevadensis A. Gray (603, 708, 772) Occasional on open slopes; 2600 - 3400 m. Lowland California.

Arctostaphylos patula E. Greene (140, 383) Common in montane chaparrals and in dry forest opening; 2400 - 3000 m. Lowland California.

Cassiope mertensiana (Bong.) Don (844) Occasional among rocks and in moist subalpine forests; 3000 - 3400 m. Circumboreal.

Kalmia polifolia Wangenh. var. microphylla (Hook.) Calder & Roy Taylor (412) Occasional in wet meadows; 2600 - 3400 m. Circumboreal.

Ledum glandulosum Nutt. (1176) Common in the wet alpine meadows and adjoining forest in the middle canyon of Deadman Creek; 2900 - 3100 m. Circumboreal.

Orthilia secunda (L.) House (828) Occasional, found only in a semi shaded ephemeral watercourse in the red fir forest; 2700 m. Circumboreal.

Phyllodoce breweri (A. Gray) Maxim. (806, 846) Occasional among rocks near streams and in mountain hemlock forests; 3000 - 3400 m. Circumboreal. 117

Pterospora andromedea Nutt. (135) Common in red fir and mixed Jeffrey pine forest; 2400 - 3000 m. Northern Hemisphere. ο ο Pyrola asarif lia Michaux ssp. asarίf lίa (825) Occasional, found only in a semi shaded ephemeral watercourse in the red fir forest; 2700 m. Circumboreal.

Pyrola minor L. (14) Occasional in mixed red fir forest; 2600 - 3000 m. Northern Hemisphere.

Pyrola picta Smith (134, 624) Common understory plant in the red fir forest; 2400 - 3000 m. Old Cordilleran.

FABACEAE

Astragalus kentrophyta A. Gray var. danaus (Barneby) Barneby (1044) UNCOMMON, CLAPS List 4. Component of the alpine fell fields near the summit of San Joaquin Peak; 3400 - 3536 m. Old Cordilleran.

Astragalus purshii Hook. var. lectulus (S. Watson) Μ.E. Jones (180, 577, 863) Common on exposed ridges and open slopes; 2600 - 3400 m. Old Cordilleran.

Astragalus whitneyi A. Gray var. whimeyi (131, 314, 569, 1061) Common on exposed ridges and open slopes; 2600 - 3536 m. Circumboreal.

Lupinus albicaulis Hook. (152, 171, 627) Common in mixed Jeffrey pine forest and open areas and edges of red fir forest; 2400 - 3000 m. Lowland California.

Lupinus argenteus Pursh var. argenteus (192, 210, 575, 911, 1228) Common among aspen and on exposed ridges and open slopes; 3000 - 3400 m. Great Basin.

Lupinus argenteus Pursh var. meionanthus (A. Gray) Barneby (125, 958) Common in dry, open forests; 2600 - 3400 m. Great Basin.

Lupinus argenteus Pursh var. Palmeri (S. Watson) Barneby (325, 430, 783) Common on exposed ridges and open slopes; 2600 - 3000 m. Great Basin.

Lupinus breweri A. Gray var. grandiflοrus C.P. Smith (475) 2400 - 2600 m. Occasional in lodgepole pine forest on southern edge of study area; Lowland California.

Lupinus excubitus Μ.E. Jones var. excubitus (960) Common on exposed ridges and open slopes near Deadman Pass, has a distinctive and pleasant "grape soda" smell; 3000 - 3300 m. Lowland California.

Lupinus duranii Eastw. (258) RARE CLAPS List 1B. A dominant plant in the Parma rabbitbrush scrub and an understory component in the Jeffrey pine forest along the trail to Glass Creek Meadow and on the south side of Glass Creek at the foot of White Wing Mountain; 2620 - 2700 m. Great Basin. 118

Lupinus lepidus Douglas var. lobbii (S. Watson) C. Hitchc. (91, 451, 502, 611, 718, 961, 726) Common in dry meadows, mixed forests, and on open pumice, slopes, and ridges; 2400 - 3400 m. Old Cordilleran.

Lupinus lepidus Douglas var. ramosus Jepson (791, 1040) Common, localized near the summit of San Joaquin Peak; 3400 - 3536 m. Old Cordilleran.

Lupinus lepidus Douglas var. sellulus (Kellogg) Barneby (274, 1002) Common in mixed and red fir forests; 2600 - 3000 m. Old Cordilleran.

Lupinus polyphyllus Lindley var. burkei (S. Watson) C. Hitchc. (13, 781, 849, 650) Common along streams and in wet meadows; 2400 - 3400 m. Circumboreal.

Trifolium cyathiferum Lindley (939) Common along streams in the red for forest; 2600 - 3000 m. Lowland California.

Trifolium longipes Nutt. var. nevadense Jepson (65, 110, 265, 1109) Abundant in wet meadows; 2600 - 3000 m. Old Cordilleran.

Trifolium monanthum A. Gray var. monanthum (35, 679, 747) Common along streams and in wet meadows; 2600 - 3400 m. Old Cordilleran.

FAGACEAE

Chrvsolepis sempervirens (Kellogg) Hjelmq. (141) Common in montane chaparrals and in mixed Jeffrey pine forest; 2400 - 3000 m. Old Cordilleran.

Quercus vacciniifolia Kellogg (139) Common in montane chaparrals and in mixed Jeffrey pine forest; 2400 - 3000 m. Lowland California.

GENTIANACEAE

Gentianopsis simplex (A. Gray) Iltis. (235, 288) Occasional in wet meadows; 2600 - 3000 m. Old Cordilleran.

Swertia radiata (Kellogg) Kuntze (1181) Occasional in the forest edges and open rocky sites of the limestone area; 2900 - 3040 m. Old Cordilleran.

GROSSULARIACEAE

Ribes cereum Douglas var. cereum (144, 380, 384, 472) Common in dry forests, montane chaparrals, and on exposed ridges and peaks; 2400 - 3536 m. Old Cordilleran.

Ribes inerme Rydb. var. inerme (89, 262) Occasional around the bases of willow clumps in Glass Creek Meadow; 2600 - 3000 m. Old Cordilleran.

Ribes montigenum McClatchie (9, 250, 365, 419, 600, 826) Common in forest Openings and in rocks on exposed ridges; 2400 - 3400 m. Old Cordilleran. 119

HYDROPHYLLACEAE

Nernophila spathulata Coy. (121, 886, 677) Common in wet meadows; 2600 - 3000 m. Lowland California.

Phacelia hastata Lehm. ssp. compacta (Brand) Heckard (53, 186, 277, 295, 338, 393, 1034) Common in dry areas throughout; 2600 - 3536 m. Great Basin.

Phacelia mutabilis E. Greene (631, 821, 938) Occasional among sagebrush and aspen; 2600 - 3000 m. Lowland California.

Phacelia ramosissima Lehm. var. ramosissima (910, 661) Occasional in montane chaparral; 2600 - 3000 m. Lowland California.

LAMIACEAE

Agastaches urticifolia (Benth.) Kuntze (630) Occasional among aspen, willows, and sagebrush in the avalanche zone along Deadman Creek. 2600 - 2700 m. Old Cordilleran.

Monardella glauca E. Greene (305, 349, 1190, 1225) Occasional on open rocky areas including the limestone outcrop; 2900 - 3040 m. Lowland California.

Monardella odoratissima Benth. ssp. odoratissima (660, 665, 695, 782, 982, 1004) Common in rocky areas among montane chaparral and in forest openings; 2400 - 3000 m. Lowland California.

Monardella odoratissima Benth. ssp. pallida (A.A. Heller) Epling (298) Occasional in rocky areas among montane chaparral; 2600 - 3000 m. Lowland California.

LINACEAE

Linum lewisii Pursh ssp. lewisii (363, 1195) Occasional on exposed ridges. 2600 - 3400 m. Old Cordilleran.

LOASACEAE

Mentzelia albicaulis Hook. (193) Occasional on pumice slopes; 3000 - 3400 m. Great Basin.

Mentzelia laevicaulis (Hook.) Torrey & Α. Gray (194) Occasional on pumice slopes; 3000 - 3400 m. Great Basin.

Mentzelia montana (Davidson) Davidson (398, 448, 573, 858) Common in open pumice and on exposed ridges and slopes; 2600 - 3400 m. Great Basin. 120

ONAGRACEAE Epilobium anagallidίfolium Lam. (1143, 1150) Occasional in high elevation riparian areas, found only along creek just below San Joaquin bowl; 3000 - 3400 m. Circumboreal.

Epilobium angustlfolium L. ssp. circumvagum Mosq. (268, 619) Common along streams; 2400 - 3000 m. Circumboreal.

Epilobium ciliatum Raf. ssp. ciliatum (86, 160, 248, 346, 563, 923, 997) Common in wet meadows and along streams; 2400 - 3400 m. Circumboreal.

Epilobium ciliatum Raf. ssp. glandulosum-(Lehm.) P. Hoch & Raven (647) Occasional in wet meadows; 2600 - 3000 m. Circumboreal.

Epilobium glaberrimum Barbey ssp. fastigiatum (Nutt.) P. Hoch & Raven (557, 653, 1013) Common in wet meadows and among aspen; 2600 - 3000 m. Circumboreal.

Epilobium glaberrimum Barbey ssp. glaberrimum (618) Occasional in dry meadows; 2400 - 2600 m. Circumboreal.

Epilobium hornemannii Reichb. ssp. hornemannii (18, 26, 780) Common along streams; 2400 - 3000 m. Circumboreal.

Epilobium lactiflorum Hausskn. (512) Occasional in wet meadows and riparian areas; 2600 - 3000 m. Circumboreal.

Epilobium obcordatum A. Gray (297, 807) Occasional in high elevation moist places; 3000 - 3400 m. Circumboreal.

Epilobium oregonense Hausskn. (525, 857, 943, 948, 1031, 1173) Common in wet meadows and riparian areas; 2600 - 3400 m. Circumboreal.

Epilobium saximontanum Hausskn. (6, 118, 486, 728, 1094, 1136) Common in wet meadows and riparian areas; 2400 - 3400 m. Circumboreal. ν Gayophytum diffusum Torrey & A. Gray ssp. par ίflorum Harlan Lewis & J. Szweykowski (46, 52, 67, 79, 163, 256, 450, 464, 511, 1035, 1082, 1137, 1153, 1215) Abundant in dry areas throughout; 2400 - 3400 m. Old Cordilleran.

OROBANCHACEAE

Orobanche corymbosa (Rydb.) Ferris (1068, 1069) Occasional among sagebrush; 2600 - 3000 m. Great Basin.

Orobanche fascίculata Nutt. (582, 1070) Occasional among sagebrush; 2600 - 3400 m. Circumboreal. 121

Orobanche uniflora L. (1177) Occasional among sagebrush, found only near Deadman Creek in the avalanche zone; 2670 m. Northern Hemisphere.

POLEΜΟΝIΑCEΑΕ

Collomia linearis Nutt. (114, 309, 678, 719) Common in dry meadows and montane chaparral; 2600 -3000 m. Old Cordilleran.

Collomia tinctoria Kellogg (583, 682, 722, 1121) Common on open slopes and in montane chaparral; 2600 - 3400 m. Old Cordilleran.

Gilia inconspicua (Smith) Sweet (860) Occasional on exposed ridges, found only on the bench between the Glass and Deadman drainages; 2600 - 3000 m. Lowland California.

Ipomopsis aggregate (Pursh) V. Grant ssp. formosissima (E. Greene) Wherry (216, 684, 909) Common among aspen, montane chaparral, on exposed ridges and in avalanche areas; 2600 - 3000 m. Lowland California.

Leptodactylon pun gens (Torrey) Rydb. (40) Common on open slopes and exposed ridges; 2600 - 3536 m. Lowland California.

Linanthus ciliatus (Benth.) E. Greene (170, 385, 439) Occasional in mixed Jeffrey pine forest; 2400 - 2600 m. Lowland California.

Linanthus nuttallii (Α. Gray) Milliken ssp. pubescens R. Patterson (621) Common in the lower, dry meadow portion of Crater Flat; 2400 - 2600 m. Lowland California.

Phlox condensata (Α. Gray) E. Nelson (1048) Common component of the alpine fell fields near the summit of San Joaquin Peak; 3400 - 3536 m. Old Cordilleran.

Phlox diffuse Benth. (587, 604, 687, 865, 1187) Common in forest openings, on open slopes and exposed ridges, and on the limestone; 2600 - 3400 m. Old Cordilleran.

Phlox gracilis E. Greene (466, 607) Common in dry meadows; 2400 - 2600 m. Old Cordilleran.

Phlox stansburyi (Torrey) A.A. Heller (688) Occasional in openings in the upper red fir forest; 2600 - 3000 m. Old Cordilleran.

Polemonium occidentale E. Greene (81, 560, 758) Occasional in wet meadows. 2600 - 3000 m. Old Cordilleran.

POLYGONACEAE

Eriogonum esmeraldense S. Watson var. esmeraldense (479) Common in the Jeffrey pine forest; 2400 - 2600 m. Great Basin. 122

Eriogonum incanum Torrey & A. Gray (207, 327, 329) Common on open slopes, bowls, and exposed ridges; 3000 - 3400 m. Old Cordilleran.

Eriogonum lobbii Torrey & A. Gray var. lobbii (199) Occasional on open slopes and among whitebark pine krummholz; 2600 - 3536 m. Great Basin.

Eriogonum microthecum Nutt. var. alpinum Rev. (317, 714, 1073). Uncommon (Hickman 1993). Common on the bench between the Glass and Deadman drainages, on metamorphic rock in the alpine, and in open pumice with Lupinus duranii; 2600 - 3536 m. Great Basin.

Eriogonum microthecum Nutt. var. laxiflorum Hook. (311, 699, 704, 705, 761, 766) Common in subalpine forests on open slopes and exposed ridges; 2600 - 3536 m. Great Basin.

Eriogonum nudum Benth. var. deductum (E. Greene) Jepson (126, 148, 204, 299) Common in dry forests and montane chaparral; 2400 - 3400 m. Lowland California.

Eriogonum nudum Benth. var. nudum (666, 1001) Common in open sagebrush; 2400 - 3000 m. Lowland California.

Eriogonum ονalifοlium Nutt. var. nivale (Canby) Μ.E. Jones (49, 181, 794) Common in open pumice and on exposed ridges and peaks; 2600 - 3536 m. Old Cordilleran.

Eriogonum rosense Nelson & Kenn. (1062, 1085, 1168). Uncommon (Hickman 1993). Common on alpine ridges below San Joaquin Peak and in open pumice with Lupinus duranii ; 2600 - 3536 m. Great Basin.

Eriogonum speruglinum A. Gray var. reddingianum (Μ.E. Jones) J. Howell (172, 315, 426, 461, 546, 579) Common in dry forests and on exposed ridges; 2400 - 3400 m. Lowland California.

Eriogonum strictum Benth. var. anserinum (Ε. Greene) R. Davis (341, 453, 455) Occasional in open pumice and on exposed ridges; 2400 - 3400 m. Old Cordilleran.

Eriogonum umbellatum Torrey var. nevadense Grand. (45, 183, 340, 617) Common in dry forests, montane chaparral, and open pumice and on exposed ridges; 2400 - 3400 m. Old Cordilleran.

Oxyria dignya (L. ) Hill (352, 789) Occasional in shaded rocks and on exposed ridges and peaks; 2600 - 3536 m. Circumboreal.

Polygonum arenastrum Boreau (936) Common in wet meadows, widespread throughout Glass Creek Meadow; 2600 - 3000 m. Alien, introduced from Europe.

Polygonum douglasii E. Greene ssp. douglasii (109, 526, 739) Common in wet meadows; 2600 - 3000 m. Old Cordilleran. 123

Polygonum douglasii E. Greene ssp. johnstonii (Munz) J. Hickman (655) Common in riparian areas and in avalanche zones; 2600 - 3000 m. Old Cordilleran.

Polygonum polygaliodes Meissner ssp. kelloggii (E. Greene) J. Hickman (523, 752, 1032) Common in wet meadows; 2600 - 3000 m. Old Cordilleran.

Polygonum shastense A. Gray (595, 800) Occasional in rocks on exposed ridges; 3000 - 3400 m. Old Cordilleran.

Rumex paucifolius S. Watson (60, 343, 407, 446, 473, 513) Common in dry meadows and forests and on open slopes and exposed ridges; 2400 - 3400 m. Old Cordilleran.

Rumex salicifolius J.A. Weinm. var. denticulatus Torrey (36, 238, 281) Common in meadows; 2400 - 3000 m. Old Cordilleran. PORTULACACEAE

Calyptridium monospermum E. Greene (889) Common in dry areas in Minaret Meadow; 2600 - 3000 m. Old Cordilleran.

Calyptridium umbellatum (Torrey) E. Greene (42, 390) Common in open pumice, dry meadow edges, and on exposed ridges; 2400 - 3000 m. Old Cordilleran.

Calyptridium umbellatum (Torrey) E. Greene) var. caudiciferum(A. Gray) Jepson (205) Common on open slopes and exposed ridges; 3000 - 3536 m. Old Cordilleran.

Lewisia nevadensis (A. Gray) Robinson (535, 883) Occasional in wet meadows; 2600 - 3000 m. Old Cordilleran.

Lewisia pygmaea (A. Gray) Robinson (1141, 1159) Occasional in high elevation moist areas, found only along stream below San Joaquin bowl; 3000 - 3400 m. Old Cordilleran.

Montia chamissoi (Sprengel) E. Greene (85, 116, 499, 510, 654) Common in wet meadows; 2400 - 3000 m. Circumboreal. PRIMULACEAE

Dodecatheon alpinum (A. Gray) E. Greene (1200) Occasional in wet alpine meadows in the middle canyon of Deadman Creek; 3000 - 3100 m. Old Cordilleran.

Primula suffructescens A. Gray (362, 805) Occasional among rocks, on open slopes, and in mountain hemlock forests; 3000 - 3536 m. Old Cordilleran. 124

RANUNCULACEAE

Aconitum columbianum Nutt. (759) Common along streams and in wet meadows; 2600 - 3000 m. Circumboreal.

Actaea rubra (Aiton) Willd. (824) Occasional in moist openings in the red for forest; 2600 - 3000 m. Northern Hemisphere.

Anemone occidentalis S. Watson (597, 605) Occasional on open slopes; 3000 - 3400 m. Old Cordilleran.

Aquilegia formosa Fischer (469) Common along streams and in wet meadows; 2400 - 3400 m. Old Cordilleran.

Delphinium depaupercrtum Torrey & A. Gray (721, 723, 754) Occasional in moist forest openings and in wet meadows; 2600 - 3000 m. Old Cordilleran.

Delphinium glaucum S. Watson (287, 622, 651) Common in wet meadows and riparian areas; 2600 - 3000 m. Old Cordilleran.

Delphinium gracilentum E. Greene (937, 1106). Range extension to the east. One small population among the willows in Minaret Meadow; 2690 m. Old Cordilleran.

Delphinium polycladon Eastw. (242, 724, 741, 977, 979, 1020, 1098) Common in wet meadows and along streams; 2600 - 3000 m. Old Cordilleran.

Ranunculus alismifolius Benth. var. alismellus A. Gray (405, 866) Common in wet meadows; 2600 - 3000 m. Lowland California.

Ranunculus alismifolius Benth. var. alismifolius (59, 379, 404) Abundant in wet meadows, one of the dominant plants in Glass Creek Meadow. 2600 - 3000 m. Lowland California.

Ranunculus eschscholzii Schldl. var. oxynotus (A. Gray) Jepson (359, 564, 1125, 1229) Occasional on open slopes and exposed ridges; 3000 - 3536 m. Circumboreal.

Ranunculus glaberrimus Hook. var. ellipticus (E. Greene) E. Greene (891) Occasional in wet meadows, found only in Minaret Meadow; 2600 - 3000 m. Old Cordilleran.

Thalictrum fendleri A. Gray var. fendleri (28, 263, 552, 639, 1010) Common along streams and in wet meadows; 2400 - 3000 m. Old Cordilleran.

RHAMNACEAE

Ceanothus velutinus Hook. var. velutinus (147, 427) Abundant in montane chaparral and in mixed Jeffrey pine forest; 2400 - 3000 m. Lowland California. 125

ROSACEAS

Amelanchier utahensis Koehne (868) Occasional in montane chaparral; 2600 - 3000 m. Great Basin.

Cercocarpus ledifolius Nutt. var. intermontanus N. Holmgren (425) Occasional, found only in the Jeffrey pine forest on the south side of Obsidian Dome; 2400 - 2420 m. Great Basin.

Chamaebatiaria millefolium (Torrey) Maxim. (480) Occasional on rocks, found only in the Jeffrey pine forest on the south side of Obsidian Dome; 2400 - 2420 m. Old Cordilleran.

Geum macrophyllum Willd. (12, 880) Occasional in wet meadows; 2600 - 3000 m. Circumboreal.

Holodiscus microphyllus Rydb. var. microphyllus (21, 146, 477, 519, 700) Common in montane chaparral, on rocks, and in mixed Jeffrey pine forest; 2400 - 3000 m. Lowland California. ρ ο Horkelia fusca Lindley ssp. arvίfl ra (Nutt.) Keck (103, 928, 929) Common in dry meadow edges; 2600 - 3000 m. Lowland California.

Ivesia lycopodioides A. Gray ssp. lycopodioides (790) Occasional in the alpine fell fields near the summit of San Joaquin Peak; 3400 - 3536 m. Lowland California.

Potentilla drummondii Lehm. ssp. breweri (S. Watson) B. Enter (1140) Occasional in high elevation riparian areas, found only along stream in San Joaquin bowl; 3000 - 3400 m. Circumboreal.

Potentilla flabellifolia Hook. (799, 1172) Occasional in high elevation riparian areas, found only along stream in San Joaquin bowl; 3000 - 3400 m. Circumboreal.

Potentilla glandulosa Lindley ssp. nevadensis (S. Watson) Keck (301, 467, 517, 601, 614, 632, 934, 1116) Common in wet meadows and along streams; 2400 - 3000 m. Lowland California.

Potentilla gracilis Hook. ssp. fastigiate (Nutt.) S. Watson (62, 278, 608, 612) Occasional in dry meadows; 2400 - 3000 m. Circumboreal.

Potentilla grayi S. Watson (1196, 1223) Occasional in high elevation wet meadows; 3000 - 3400 m. Old Cordilleran.

Prunus emarginata (Hook.) Walp. (217, 386, 667, 972) Common in montane chaparral, among rocks, and along streams; 2400 - 3000 m. Old Cordilleran.

Purshia tridentata (Pursh) DC var. glandulosa (Curran) Μ.E. Jones (395, 710) Common understory component of the Jeffrey and Limber pine forests on the upper slopes of North Point; 2950 - 2987 m. Great Basin. 126

Purshia tridentata (Pursh) DC var. tridentata (127, 663) Common component of the montane chaparrals, dominant on mid-elevation exposed ridges; 2400 - 3000 m. Great Basin.

Sibbaldia procumbens L. (1127) One small population near a rocky spring along the creek just below San Joaquin bowl; 3180 m. Circumboreal.

Sorbus californίca E. Greene (779, 848) Occasional in riparian areas only in the south canyon of Deadman Creek; 2700 - 2800 m. Circumboreal.

RUBIACEAE

Galium bifolium S. Watson (634) Occasional in open sagebrush and willows in avalanche areas; 2600 - 3000 m. Old Cordilleran.

Galium hypotrichium A. Gray ssp. hypotrichium (516, 701, 962) Occasional in rocks in exposed sites; 2600 - 3536 m. Old Cordilleran.

Galium trifidum L. var. pacificum Wieg. (551) Occasional in rocks in mixed forests; 2600 - 3000 m. Lowland California.

Galium triflorum Michaux (827, 940) Occasional in moist areas in the red fir forest; 2600 - 3000 m. Northern Hemisphere.

Kelloggia galioides Torrey (549) Occasional in rocks in mixed forests; 2600 - 3000 m. Old Cordilleran.

SALICACEAE

Populus balsamifera L. ssp. trichocarpa (Torrey & A. Gray) Brayshaw (855) A single tree next to the now closed upper section of Deadman Road; 2580m. Lowland California.

Populus tremuloides Michaux (214) Abundant on moist steep slopes, in avalanche zones, and in moist forest openings; 2600 - 3000 m. Old Cordilleran.

Salix arctica Pallas (1212) Occasional in wet alpine meadows in the middle canyon of Deadman Creek; 3000 - 3100 m. Old Cordilleran.

Salix eastwoodiae A.A. Heller (99, 100, 271, 272, 403, 414, 415, 534) Common in wet meadows; 2600 - 3000 m. Old Cordilleran.

Salix geyeriana Andersson (259, 273, 353, 371, 416, 492, 874, 1026) Common in wet meadows, along streams, and in moist forest openings; 2400 - 3400 m. Old Cordilleran.

Salix jepsonii C. Schneider (387, 767) Occasional along streams; 2400 - 3000 m. Old Cordilleran. 127

Salix lasiolepis Benth. (869, 971) Occasional along streams and in moist forest openings; 2400 - 2600 m. Lowland California.

Salix lemmonii Bebb (31, 381, 382, 636, 745, 755, 890, 985, 1022) Common in wet meadows and along streams; 2400 - 3000 m. Old Cordilleran.

Salix orestera C. Schneider (266, 270, 400, 768, 778, 796, 798, 839, 843, 870, 953, 1145, 1151) Abundant in wet meadows and along streams; 2400 - 3400 m. Old Cordilleran.

SAXIFRAGACEAE

Heuchera micrantha Lindley var. erubescens (A. Braun & C. Bouch) C. Rosend. (599, 702, 916) Occasional among rocks; 2600 - 3400 m. Lowland California.

Lithophragma glabrum Nutt. (1132) One small population near a rocky spring along the creek just below San Joaquin bowl; 3180 m. Lowland California.

Mitella breweri A. Gray (850, 978) Occasional in wet meadows and along streams; 2600 - 3000 m. Circumboreal.

Saxifraga aprica E. Greene (867) Occasional in wet meadows, found only in Minaret Meadow; 2600 - 3400 m. Old Cordilleran.

Saxifrage odontoloma Piper (19) Common in wet meadows and along streams; 2400 - 3400 m. Old Cordilleran.

Saxifrage tolmiei Torrey & A. Gray (345) Occasional in rocks on exposed ridges; 3000 - 3400 m. Old Cordilleran.

SCROPHULARIACEAE

Castilleja applegatei Fern. ssp. disticha (Eastw.) Chuang & Heckard (956) Occasional on exposed ridges; 3000 - 3400 m. Old Cordilleran.

Castilleja applegatei Fern. ssp. pallida (Eastw.) Chuang & Heckard (1226) Occasional on the limestone outcrop; 2900-3040 m. Old Cordilleran.

Castilleja applegatei Fern. ssp. pinetorum (Fern.) Chuang & Heckard (128, 185, 347) Common on open slopes and exposed ridges; 2400 - 3400 m. Old Cordilleran.

Castilleja lemmonii A. Gray (922) One small population in the middle of Glass Creek Meadow; 2600 - 3000 m. Old Cordilleran.

Castilleja linariifοlia Benth. (227, 318, 764, 899) Uncommon high elevation pink color morph (Howald 1983). Common among rocks, sagebrush, and montane chaparral on exposed slopes and ridges, especially on the bench between the Glass and Deadman watersheds; 2600 - 3000 m. Great Basin. 128

Castilleja miniata Hook. ssp. miniata (16, 773) Common in wet meadows and along streams; 2400 - 3400 m. Old Cordilleran.

Castilleja nana Eastw. (211, 333) Occasional on exposed ridges; 3000 - 3536m. Old Cordilleran.

Collinsia parviflora Lindley (120, 508, 656, 727, 861) Common in wet meadows and on exposed mid elevation ridges; 2600 - 3000 m. Old Cordilleran.

Mimulus breweri (E. Greene) Coy. (743, 1236) Occasional in wet meadows and moist forest openings; 2600 - 3000 m. Lowland California.

Mimulus lewisii Pursh (3, 221) Common along streams and in wet meadows; 2400 - 3000 m. Old Cordilleran.

Mimulus mephiticus E. Greene (37, 188, 326, 862, 1058, 1072) Common in open pumice and on exposed ridges; 2600 - 3536 m. Lowland California.

Mimulus moschatus Lindley (1233) One small population on a sandy, steep, wet forest slope in the middle canyon of Deadman Creek; 2760 m. Old Cordilleran.

Mimulus nanus Hook. & Am. (392, 445) Common in open pumice and mixed Jeffrey pine forest; 2400 - 3000 m. Old Cordilleran.

Mimulus primuloides Benth. ssp. primuloides (61, 505, 1089) Common in wet meadows; 2600 - 3400 m. Old Cordilleran.

Mimulus suksdorfiί A. Gray (1135, 1154) Common but localized in high elevation open moist areas; 3000 - 3400 m. Old Cordilleran.

Mimulus tilingii Regel (4, 23, 58, 87, 290, 487, 514, 561, 649, 879, 1096, 1128) Common in wet meadow and along streams; 2400 - 3400 m. Old Cordilleran.

Pedicularis attollens A. Gray (63, 509) Common in wet meadows; 2600 - 3400 m. Circumboreal.

Pedicularis groenlandica Retz. (88, 507) Common in wet meadows; 2600 - 3000 m. Circumboreal.

Pedicularis semibarbata A. Gray (167, 717) Occasional in the red fir forest. 2400 - 2600 m. Lowland California.

Penstemon davidsonii E. Greene var. davidsonii (366, 1045) Occasional on open slopes and exposed ridges; 3000 - 3536 m. Old Cordilleran.

Penstemon heterodoxus A. Gray var. cephalophorus (E. Greene) N. Holmgren (74, 122, 458, 494, 503) First report for Mono County. Common on dry meadow edges and in pumice at Crater Flat; 2400 - 3000 m. Old Cordilleran. 129

Penstemon heterodoxus Α. Gray var. heterodoxus (462, 474, 593, 686, 795, 1053, 1133) Common in dry meadows, open pumice, forest openings, and on exposed ridges and peaks; 2400 - 3536 m. Old Cordilleran.

Penstemon newberryi Α. Gray var. newberryi (1186) Occasional in rocks and on the limestone; 2600 - 3400 m. Old Cordilleran.

Penstemon rostriflοrus Kellogg (124, 482) Occasional in the Jeffrey pine and mixed Jeffrey pine forests; 2400 - 2600 m. Old Cordilleran.

Penstemon rydbergii Nelson var. oreocharis (E. Greene) N. Holmgren (230, 924) Occasional in dry meadow edges, found only in Glass Creek Meadow; 2700 m. Old Cordilleran.

Penstemon speciosus Lindley (182, 318, 357, 364) Common in open pumice and on exposed ridges; 2600 - 3536 m. Old Cordilleran.

Veronica americana (Raf.) Schwein. (93, 638) Common in wet meadows and riparian areas; 2600 - 3000 m. Northern Hemisphere.

Veronica serpyllifolia L. ssp. humifusa (Dickson) Syme (108, 550, 734) Common in wet meadows, widespread in Glass Creek Meadow; 2600 - 3000 m. Alien, introduced from Europe.

Veronica wormskjoldii Roemer & Schultes (811, 830, 1146, 1161, 1204) Common in wet meadows and riparian areas; 3000 - 3400 m. Old Cordilleran.

SOLANACEAE

Chamaesaracha nana (A. Gray) A. Gray (547, 1117) Occasional in rocky montane chaparral areas; 2600 - 3000 m. Old Cordilleran.

URTICΑCEΑE

Urtica dioica L. ssp. holosericea (Nutt.) Thorne (155, 224) Occasional in moist areas; 2400 - 3000 m. Northern Hemisphere.

VALERIANACEAE

Valeriana calίfornίca A.A. Heller (306, 355, 447) Occasional in subalpine forests and on open slopes and exposed ridges; 2600 - 3400 m. Circumboreal.

VIOLACEAE

Viola macloskeyi F. Lloyd (411) Occasional in wet meadows; 2600 - 3400 m. Circumboreal.

Viola pinetorum E. Greene ssp. pinetorum (585) Occasional on open slopes; 3000 - 3400 m. Lowland California. 130

Viola purpurea Kellogg ssp. integrifοlia M. Baker & J. Clausen (438) Occasional in moist forest openings; 2600 - 3000 m. Lowland California.

MONOCOTS

CYPERACEAE

Carex albonigra Mackenzie (1051) Common in the dry sedge meadows near the summit of San Joaquin Peak; 3400 - 3536 m. Circumboreal.

Carex breweri Boott var. breweri (206, 324, 368, 804, 815) Common in open pumice and on open slopes and exposed ridges; 3000 - 3536 m. Circumboreal.

Carex douglasii Boott (112, 316, 465, 540, 609) Common in dry meadows; 2400 - 3000 m. Old Cordilleran. ο Carex filίf lίa Nutt. var. erostrata Kϋk. (919) Common in wet meadows; 2600 - 3000 m. Old Cordilleran.

Carex helleri Mackenzie (354) Common on exposed ridges and open slopes; 3000 - 3400 m. Circumboreal.

Carex heteroneura W. Boott var. epapillosa (Mackenzie) F. Herr. (786, 1147) Common on exposed ridges and open slopes; 3000 - 3536 m. Old Cordilleran.

Carex heteroneura W. Boott var. heteroneura (15, 104, 254, 344, 770, 995, 1206) Abundant in dry meadows and forests, in rocks, and on exposed ridges and open slopes; 2400 - 3400 m. Old Cordilleran.

Carex illota L. Bailey (926, 1025) Occasional in wet meadows; 2600 - 3000 m. Old Cordilleran.

Carex jonesίi L. Bailey (671, 674, 931, 944, 1199) Common in wet meadows and riparian areas; 2600 - 3400 m. Old Cordilleran.

Carex lenticularis Michaux var. lipocarpa (Holm) L. Standley (945, 1198, 1208, 1211, 1232) Common in wet meadows and riparian areas; 2600 - 3536m. Circumboreal.

Carex leporinella Mackenzie (166, 813, 819) Occasional in dry meadows and stream beds and on open slopes; 2400 - 3400 m. Old Cordilleran.

Carex microptera Mackenzie (1157) Occasional in riparian areas; 3000 - 3400m. Old Cordilleran.

Carex multicostata Mackenzie (33, 544, 586, 613, 841) Common in dry forests, meadows, and riparian areas and on open slopes; 2400 - 3400 m. Old Cordilleran. 131

Carex nehrascensis Dewey (64, 71, 676) Common in wet meadows, especially in saturated areas; 2600 - 3000 m. Old Cordilleran.

Carex nervina L. Bailey (537, 984, 1014) Common in wet meadows and spring areas; 2400 - 3000 m. Lowland California.

Carex nigricans C. Meyer (1170) Occasional on open slopes and exposed ridges; 3000 - 3536 m. Old Cordilleran.

Carex petasata Dewey (774) Rare in California, CLAPS List 2. One population in a seep on the north side of the south canyon of Deadman Creek; 2800 m. Old Cordilleran.

Carex phaeocephala Piper (1139) Occasional in high elevation moist areas; 3000 - 3400 m. Old Cordilleran.

Carex praegracilis W. Boott (531) Occasional in wet meadows; 2600 - 3000 m. Lowland California.

Carex praticola Rydb. (1049) Rare in California, CLAPS List 2. Common in the dry sedge meadows near the summit of San Joaquin Peak; 3400 - 3536 m. Old Cordilleran.

Carex preslii Steudel (729) Occasional in forests and dry stream beds; 2600 - 3000 m. Old Cordilleran.

Carex propositi Mackenzie (406) Uncommon (Hickman 1993). Occasional in wet meadows; 2600 - 3000 m. Old Cordilleran.

Carex rossii Boott (872, 893) Common in lodgepole and Jeffrey pine forests; 2400 - 2600 m. Old Cordilleran.

Carex specifica L. Bailey (838, 852) Occasional in riparian areas, found only in the south canyon of Deadman Creek; 2600 - 3000 m. Lowland California.

Carex spectabilis Dewey (836, 847, 994, 1163, 1167) Common in riparian areas; 2400 - 3400 m. Old Cordilleran.

Carex stramίniformis L. Bailey (350, 615) Common in dry riparian areas and on exposed ridges and open slopes; 2400 - 3400 m. Old California.

Carex subnigricans Stacey (1217) Occasional in wet meadows; 2600 - 3000 m. Old Cordilleran.

Carex tahoensis F.J. Smiley (1046, 1064, 1235) Uncommon (Hickman 1993) Common on San Joaquin Ridge and near the summit of San Joaquin Peak; 3000 - 3536 m. Lowland California.

Carex utriculata Boott (98, 286) Common in wet meadows; 2600 - 3400 m. Northern Hemisphere. 132

Carex vesicaria L. var. vesicaria (749) Occasional in wet meadows; 2600 - 3000 m. Northern Hemisphere.

Eleocharis acicularis (L.) Roemer & Schultes var. bella Piper (527) Occasional in wet meadows; 2600 - 3000 m. Northern Hemisphere.

Eleocharis parvula (Roemer & Schultes) Link (930) UNCOMMON, CLAPS List 4. One population in the north-west center of Glass Creek Meadow; 2680 m. Circumboreal.

IRIDACEAE

Iris missouriensis Nutt. (82) Occasional in Glass Creek Meadow; 2680 m. Old Cordilleran.

JUNCACEAE

Juncus chlorocephalus Engelm. (539, 748) Common in meadows; 2600 - 3000 m. Lowland California.

Juncus drummondii E. Meyer (237, 831) Common in meadows and riparian areas; 2600 - 3536 m. Old Cordilleran.

Juncus longistylis Torrey (648) Occasional in riparian areas; 2600 - 3000 m. Northern Hemisphere.

Juncus mertensianus Bong. (29,30, 78, 115, 413, 504, 1188, 1210) Abundant in wet meadows and riparian areas; 2600 - 3400 m. Circumboreal.

Juncus mexicanus Willd. (107, 495, 496, 521, 1091) Common in wet meadows; 2400 - 3000 m. Lowland California.

Juncus nevadensis S. Watson (620, 885, 892, 969, 986, 1023, 1092) Common in dry meadows, forests, and riparian areas; 2400 - 3000 m. Old Cordilleran.

Juncus orthophyllus Coy. (77, 522) Common in wet meadows; 2600 - 3000 m. Old Cordilleran.

Juncus parryi Engelm. (189, 342, 423, 812) Common on dry slopes and open bowls, in rocks and on exposed ridges; 2400 - 3536 m. Old Cordilleran.

Luzula comosa E. Meyer (538, 1050, 1165) Occasional in meadows and on open slopes and exposed ridges; 2600 - 3536 m. Old Cordilleran.

Luzula divaricata S. Watson (592) Occasional on open scree slopes; 3000 - 3536 m. Lowland California.

Luzula orestera Sharsm. (105, 884) Occasional in wet meadows; 2600 - 3000 m. Old Cordilleran. 133

Luzula parviflora (Ehrh.) Desv. (11, 245, 488, 993) Occasional in wet meadows and riparian areas; 2400 - 3000 m. Northern Hemisphere.

Luzula spicata (L.) DC (1171) Occasional in red fir/mountain hemlock stands, subalpine forest and on surrounding slopes; 3000 - 3400 m. Circumboreal.

Luzula subcongesta (S. Watson) Jepson (1142, 1205) Occasional in high elevation riparian areas; 3000 - 3400 m. Lowland California.

LILIACEAE

Album bisceptrum S. Watson var. bisceptrum (633) Occasional in sagebrush, aspen, and willows in avalanche areas; 2600 - 3000 m. Great Basin.

Allium campanulatum S. Watson (643, 644, 662, 672, 725, 730, 732, 932, 949, 1019) Common in meadows and riparian areas; 2600 - 3000 m. Old Cordilleran.

Allium obtusum Lemmon var. obtusum (594, 1156) Occasional on open slopes and exposed ridges in metamorphic soil; 3000 - 3400 m. Old Cordilleran.

Allium validum S. Watson (498) Common in wet meadows and along streams; 2400 - 3400 m. Old Cordilleran.

Calochortus bruneaunis Nelson & J.F. Macbr. (454, 659, 901, 1017) Common in open sagebrush and on mid elevation exposed ridges; 2400 - 3000 m. Old Cordilleran.

Fritillaria pinetorum A. Davids (584, 859) Common, localized on the bench between the Glass and Deadman watersheds and along the stream above and to the south; 2600 - 3400 m. Lowland California.

Lilium kelleyanum Lemmon (964) Occasional near streams and springs in the red fir forest; 2400 - 2600 m. Lowland California.

Smilacina stellata (L.) Desf. (437, 873) Occasional in moist areas in the forests; 2400 - 3000 m. Circumboreal.

Veratrum californicum Durand var. californicum (236) Occasional in wet meadows, dominant in areas of upper Glass Creek Meadow; 2600 - 3000 m. Circumboreal.

ORCHIDACEAE

Platanthera hyperborea (L.) Lindley (980) Occasional along stream banks; 2600 - 3000 m. Circumboreal.

Platanthera leucostachys Lindley (616, 657, 951, 1191) Occasional in wet meadows and riparian areas; 2400 - 3400 m. Circumboreal. 134

Spiranthes romanzoffiana Cham. (232) Occasional in wet meadows. 2600 - 3000 m. Circumboreal.

POACEAE

Achnatherum hymenoides (Roemer & Schultes) Barkworth (576) Occasional on exposed ridges; 3000 - 3400 m. Great Basin.

Achnatherum lettermani (Vasey) Barkworth (73, 95) Occasional in dry meadows; 2600 - 3000 m. Old Cordilleran.

Achnatherum nelsonii (Scribner) Barkworth ssp. dorei (Barkworth & J. Maze) Barkworth (1102) Occasional in dry meadows; 2600 - 3000 m. Circumboreal.

Achnatherum nevadense (B. Johnson) Barkworth (313) Occasional on exposed ridges, found only on bench between Glass and Deadman watersheds; 2600 - 3000 m. Circumboreal.

Achnatherum occidentale (Thurber) Barkworth ssp. cal~fοrnicum (Merr. & Butt Davy) Barkworth (179, 376, 456, 856) Common among aspen and in dry forests and openings; 2400 - 3000 m. Old Cordilleran.

Achnatherum occidentale (Thurber) Barkworth ssp. occidentale (485, 914, 1166) Common in open pumice and on exposed ridges; 2400 - 3400 m. Old Cordilleran.

Agrostis exarata Trin. (34, 757) Occasional in riparian areas; 2400 - 3000 m. Lowland California.

Agrostis idahoensis Nash (25, 75, 253, 282, 554, 775, 1090) Common in meadows and riparian areas; 2400 - 3000 m. Old Cordilleran.

Agrostis pollens Trin. (738) Occasional in wet meadows; 2600 - 3000 m. Circumboreal.

Agrostis scabra Willd. (219, 276) Common in meadows and forests; 2600 - 3000 m. Circumboreal.

Agrostis thurberiana A. Hitchc. (246, 533, 840, 1209) Common in wet meadows and riparian areas; 2600 - 3536 m. Old Cordilleran.

Agrostis variabilis Rydb. (920, 1099, 1149, 1220) Common in wet meadows and riparian areas; 2600 - 3400 m. Old Cordilleran.

Alopecurus aequalis Sobol. (279) Occasional in wet meadows; 2600 - 3000 m. Circumboreal.

Bromus carinatus Hook. & Am. var. carinatus (337, 541, 565) Occasional on exposed ridges and in dry forest openings; 2600 - 3400 m. Old Cordilleran. 135

Bromus ciliatus L. (373, 820, 942) Occasional in riparian areas; 2400 - 3000 m. Circumboreal.

Bromus laevipes Shear (436) Occasional in moist forest openings; 2600 - 3000 m. Lowland California.

Bromus suksdοrfii Vasey (228, 308, 335, 996) Common in meadows and riparian areas and on exposed ridges; 2400 - 3400 m. Lowland California.

Bromus tectorum L. (864) One small population on the south-facing upper slope of the bench between the Glass and Deadman watersheds; 2600 - 3000 m. Alien, native to Eurasia.

Calamagrostis canadensis (Michaux) Beauv. (32, 267, 829, 992, 1095) Common in meadows and riparian areas; 2400 - 3000 m. Northern Hemisphere.

Calamagrostis purpurascens R.Br. (591) Occasional on open slopes; 3000 - 3400 m. Circumboreal.

Calamagrostis stricta (Timm) Koeler ssp. stricta (269, 275) Occasional in meadows; 2600 - 3000 m. Northern Hemisphere.

Cinna latifolia (Goeppert) Griseb. (161, 372) Occasional in meadows and riparian areas; 2400 - 3400 m. Northern Hemisphere.

Deschampsia elongata (Hook.) Benth. (72, 736, 925, 933, 1029, 1100) Common in meadows; 2600- 3000 m. Lowland California.

Elymus elymoides (Raf.) Swezey ssp. califοrnicus (J.G. Smith) Barkworth (39, 44, 56, 190, 332) Common in open pumice and on exposed ridges and slopes; 2600 - 3400 m. Old Cordilleran.

Elymus elymoides (Raf.) Swezey ssp. elymoides (176, 195, 302, 715) Common in dry sites throughout; 2400 - 3400 m. Old Cordilleran.

Elymus glaucus Buckley ssp. glaucus (652) Common in open sagebrush; 2600 - 3000 m. Lowland California.

Elymus sierrus Gould (814, 1122, 1182) Occasional on open slopes and exposed ridges and in rocks; 3000 - 3536 m. Old Cordilleran.

Elymus trachycaulus (Link) Shinn. ssp. trachycaulus (123, 321, 491) Occasional in dry meadows and riparian areas; 2400 - 3000 m. Northern Hemisphere.

Festuca brachyphylla Schultes & Schultes f. ssp. breviculmis S. Frederiksen (788, 1047, 1126) Occasional on exposed ridges and open slopes; 3000 - 3536 m. Circumboreal.

Glyceria elata (Lam.) A. Hitchc. (94, 280, 375, 675) Common in meadows and riparian areas; 2600 - 3000 m. Northern Hemisphere. 136

Hordeum brachyantherum Nevski ssp. californicum (Covas & Stebb.) v. Bothmer, N. Jacobsen & O. Seberg (96, 489, 740, 921) Occasional in meadows and riparian areas; 2400 - 3000 m. Old Cordilleran.

Leymus cinereus (Scribner & Merr.) A. Love (917) Occasional among rocks in mixed Jeffrey pine forest; 2600 - 3000 m. Great Basin.

Melica bulbosa Geyer (897) Occasional on exposed ridges, found only on the bench between Glass and Deadman drainages; 2600 - 3000 m. Lowland California.

Melica stricta Bolander (696) Occasional on rock outcrops and slopes; 2600 - 3000 m. Old Cordilleran. ο Muhlenbergia filif rmίs (Thurber) Rydb. (524, 753, 1219) Common in meadows; 2600 - 3000 m. Old Cordilleran.

Muhlenbergia minutissima (Steudel) Swallen (562) Occasional in meadows; 2600 - 3000 m. Old Cordilleran.

Muhlenbergia richardsonis (Trin.) Rydb. (113, 229, 559, 606) Common in dry meadows; 2600 - 3000 m. Old Cordilleran.

Phleum alpinum L. (66, 111, 842) Common in wet meadows and along streams; 2600 - 3400 m. Circumboreal.

Poa bolanderi Vasey (646, 1005) Occasional in dry riparian areas; 2400- 3000 m. Lowland California.

Poa cusickii Vasey ssp. epilis (Scribner) W.A. Weber (1124, 1202) Occasional in open rocky areas, including limestone; 2800-3100 m. Old Cordilleran.

Poa fendleriana (Steudel) Vasey ssp. longiligula (Scribner & Williams) R. Soreng (1123) Occasional in riparian areas; 3000 - 3400 m. Old Cordilleran.

Poa glauca M. Vahl. ssp. rupicola (Nash) W.A. Weber (421) Occasional in Jeffrey pine forest; 2400 - 2600 m. Old Cordilleran.

Poa keckii R. Soreng (1155) Occasional on exposed ridges and open slopes; 3000 - 3536 m. Great Basin.

Poa leptocoma Trin. ssp. leptocoma (998, 1024) Occasional in meadows and riparian areas; 2400 - 3000 m. Old Cordilleran.

Poa palustris L.(1178) Occasional in riparian areas, found only at now closed road crossing of Deadman Creek; 2600 - 3000 m. Northern Hemisphere, naturalized, native to Europe.

Poa pratensis L. ssp. pratensis (941) Occasional in riparian areas, found only at now closed road crossing of Deadman Creek; 2600 - 3000 m. Northern Hemisphere, naturalized, native to Europe. 137

Poa secunda J.S. Presl ssp. secunda (877, 1174, 1197) Occasional, restricted to the limestone outcrop. 2900 - 3040 m. Old Cordilleran.

Poa wheeleri Vasey (8, 389, 410, 422, 460, 471, 566, 588, 691, 744, 797, 973) Abundant in dry areas throughout; 2400 - 3536 m. Old Cordilleran.

Trisetum spicatum L. (Richter) (27, 106, 200, 348, 816, 896, 1009, 1107, 1129) Common in meadows and dry areas throughout; 2400 - 3400 m. Circumboreal.

Trisetum wolfii Vasey (756, 912, 935, 1042) Occasional in dry meadows and on exposed ridges, slopes, and peaks; 2600 - 3536 m. Old Cordilleran. APPENDIX B. Environmental variables.

Table 16. Environmental variables used as the secondary overlay matrix in the ordinations. Area Size Codes: 1=smaller than San Joaquin Roadless Area (SJRA), 2=same size as SJRA, 3=larger than the SJRA, 4= Regional, much larger than the SJRA. Flora numbers are as follows: 1=San Joaquin Roadless Area, 2=Harvey Monroe Hall Research Natural Area, 4=White Mountains, 5=Upper Walker River Watershed — Sierra Nevada portion, 6=Sweetwater Mountains, 7=Bodie Hills, 8=Tuolumne Meadows, 9=Lassen Volcanic National Park, 10=Trinitiy Alps, 11=San Bernardino Mountains, 12=Sequoia and Kings Canyon National Parks, 13=Desolation Wilderness, 14=Mammoth Mountain.

Flora Latitude Longitude Latitude Longitude Area Elevation Highest No. of Midpoint Midpoint difference difference Site Span (m) Elevation Taxa (Location) (Location) from SJRA from SJRA Code (m) (Distance) (Distance) 1 37°42'N 119°02'W 0' 0' 2 1136 3536 446 2 37°57'Ν 119°17'W 15' 15 1 911 3838 203 4 37°34'Ν 118°12W 8' 90' 4 1965 4343 508 5 38°22'Ν 119°27'W 80' 25' 3 1415 3793 707 6 38°26'Ν 119°20'W 84' 18' 3 1181 3559 639 7 38°15'Ν 119°00'W 73' 2' 2 682 3121 329 8 37°54'N 120° 19'W 12' 117' 2 1490 3838 475 9 40°30Ν 121°25'W 288' 233' 3 1602 3188 786 10 40°54'Ν 123°02'W 312' 400' 4 1121 2745 573 11 34°11'N 116°56'W 331' 246' 4 1676 3505 796 12 36°45'Ν 118°44'W 97' 58' 4 2285 4419 881 13 38°57'Ν 120°11W 115' 109' 2 788 3044 532 14 37°41'Ν 119°02'W 1' 0' 1 931 3370 274

138 APPENDIX C. Similarity matrices.

Table 17. Sorensen's similarity matrix for the species composition of all floras. Flora numbers are as follows: 1=San Joaquin Roadless Area, 2=Harvey Monroe Hall Research Natural Area, 4=White Mountains, 5=Upper Walker River Watershed - Sierra Nevada portion, 6=Sweetwater Mountains, 7=Bodie Hills, 8=Tuolumne Meadows, 9=Lassen Volcanic National Park, 10=Trinitiy Alps, 11=San Bernardino Mountains, 12=Sequoia and Kings Canyon National Parks, 13=Desolation Wilderness, 14=Mammoth Mountain.

Floral1 2 4 5 6 7 8 9 10 11 12 13 14 1 1 2 .445 1 4 .397 .210 1 5 .518 .326 .436 1 6 .480 .281 .471 .854 1 7 .390 .188 .402 .445 .508 1 8 .559 .490 .335 .517 .443 .271 1 9 .447 .267 .237 .459 .394 .231 .473 1 10 .356 .217 .157 .323 .277 .171 .353 .490 1 11 .295 .147 .238 .355 .343 .219 .274 .371 .249 1 12 .518 .316 .347 .551 .481 .278 .584 .568 .409 .418 1 13 .553 .420 .297 .534 .453 .304 .606 .577 .457 .305 .585 1 14 .525 .335 .286 .417 .384 .351 .407 .337 .249 .241 .372 .430 1

139 140

Table 18. Sorensen's similarity matrix for percentages of taxa in each affinity category. Flora numbers are as follows: 1=San Joaquin Roadless Area, 2=Harvey Monroe Hall Research Natural Area, 4=White Mountains, 5=Upper Walker River Watershed - Sierra Nevada portion, 6=Sweetwater Mountains, 7=Bodie Hills, 8=Tuolumne Meadows, 9=Lassen Volcanic National Park, 10=Trinitiy Alps, 11=San Bernardino Mountains, 12=Sequoia and Kings Canyon National Parks, 13=Desolation Wilderness, Ι4=Mammoth Mountain.

1 Floral 2 4 5 6 7 8 9 10 11 12 13 14 1 1 2 .852 1 4 .832 .693 1 5 .896 .757 .891 1 6 .874 .735 .905 .961 1 7 .798 .659 .901 .861 .885 1 8 .916 .936 .748 .812 .790 .714 1 9 .868 .804 .759 .860 .831 .721 .856 1 10 .857 .837 .727 .798 .785 .696 .889 .896 1 11 .791 .643 .707 .817 .800 .761 .707 .820 .777 1 12 .907 .808 .796 .861 .838 .761 .872 .918 .915 .835 1 1 3 .923 .863 .787 .853 .829 .747 .927 .920 .932 .779 .938 1 14 .878 .774 .787 .882 .882 .773 .806 .876 .825 .844 .878 .865 1