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VOL. 38, NO. 4 AND VOL. 39, NO. 1 • OCTOBER 2010 AND JANUARY 2011 FREMONTIA JOURNAL OF THE NATIVE SOCIETY

USING ALPINE FLORA TO MEASURE ECOLOGICAL EFFECTS OF CLIMATE CHANGE, SERPENTINE SOILS AND , ANDVOLUME OTHER 38:4/39:1, ARTICLES OCTOBER 2010/JANUARY 2011 FREMONTIA CALIFORNIA NATIVE PLANT SOCIETY CNPS, 2707 K Street, Suite 1; Sacramento, CA 95816-5113 FREMONTIA Phone: (916) 447-CNPS (2677) Fax: (916) 447-2727 Web site: www.cnps.org Email: [email protected] VOL. 38, NO. 4, OCTOBER 2010 AND VOL 39, NO. 1, JANUARY 2011 MEMBERSHIP Membership form located on inside back cover; Copyright © 2011 dues include subscriptions to Fremontia and the CNPS Bulletin California Native Plant Society Mariposa Lily ...... $1,500 Family or Group ...... $75 Bob Hass, Editor Benefactor ...... $600 International or Library ...... $75 Patron ...... $300 Individual ...... $45 Beth Hansen-Winter, Designer Plant Lover ...... $100 Student/Retired/Limited Income . $25 Brad Jenkins, Cynthia Roye, CORPORATE/ORGANIZATIONAL and Jake Sigg, Proofreaders 10+ Employees ...... $2,500 4-6 Employees ...... $500 7-10 Employees ...... $1,000 1-3 Employees ...... $150 CALIFORNIA NATIVE PLANT SOCIETY STAFF (SACRAMENTO) CHAPTER COUNCIL Acting Exec. Director . . . Sue Britting David Magney (Chair); Larry Levine Dedicated to the Preservation of Finance & Administration Manager . (Vice Chair); Marty Foltyn (Secretary) the California Native Flora Cari Porter Alta Peak (Tulare) . . . . Joan Stewart Membership & Development Coor- Bristlecone (Inyo-Mono) ...... The California Native Plant Society dinator ...... Stacey Flowerdew Steve McLaughlin (CNPS) is a statewide nonprofit organi- Conservation Program Director . . . . . Channel Islands . . . . David Magney zation dedicated to increasing the Greg Suba Dorothy King Young (Mendocino/ understanding and appreciation of Rare Plant Botanist . . . . Aaron Sims Sonoma Coast) . . . . . Nancy Morin California’s native plants, and to pre- Vegetation Program Director . . . Julie East Bay ...... Bill J. Hunt serving them and their natural habitats Evens El Dorado ...... Susan Britting for future generations. Vegetation Ecologists . Jennifer Buck, Kern County . . . . . Dorie Giragosian CNPS carries out its mission through Kendra Sikes Los Angeles/Santa Monica Mtns . . . . Education Program Director . . . Josie science, conservation advocacy, educa- Betsey Landis Crawford Marin County . . Carolyn Longstreth tion, and horticulture at the local, state, Administrative Assistant . . . . . Marcy Milo Baker (Sonoma County) . . . . . and federal levels. It monitors rare and Millett endangered plants and habitats; acts to Liz Parsons save endangered areas through public- STAFF (AT LARGE) Mojave Desert ...... Tim Thomas ity, persuasion, and on occasion, legal Fremontia Editor ...... Bob Hass Monterey Bay . . . . Rosemary Foster Mount Lassen ...... Catie Bishop action; provides expert testimony to CNPS Bulletin Editor . . . . . Bob Hass Napa Valley ...... Gerald Tomboc government bodies; supports the estab- Legislative Consultant .Vern Goehring East Bay Conservation Analyst . . . . . North Coast ...... Larry Levine lishment of native plant preserves; spon- North San Joaquin . . . . Alan Miller Mack Casterman sors workdays to remove invasive plants; Orange County . . . . . Nancy Heuler Website Coordinator . . Mark Naftzger and offers a range of educational activi- Redbud (Grass Valley/Auburn) . . . . ties including speaker programs, field PROGRAM ADVISORS Joan A. Jernegan trips, native plant sales, horticultural Rare Plant Program Senior Advisor . . . Riverside/San Bernardino counties . . workshops, and demonstration gardens. Jim Andre Katie Barrows Since its founding in 1965, the tradi- Sacramento Valley . . . Glen Holstein Vegetation Program Senior Advisor . . San Diego ...... David Varner tional strength of CNPS has been its Todd Keeler-Wolf San Gabriel Mtns . . . . Orchid Black dedicated volunteers. CNPS activities Horticulture Committee Chair ...... San Luis Obispo . . . . Kristie Haydu are organized at the local chapter level Laura Camp where members’ varied interests influ- Sanhedrin (Ukiah) ...... Geri CNPS Press Co-Directors . . . . . Holly Hulse-Stephens ence what is done. Volunteers from the Forbes, Dore Brown 33 CNPS chapters annually contribute Santa Clara Valley . . . Judy Fenerty Poster Program . . . Bertha McKinley, Santa Cruz County . Deanna Giuliano in excess of 97,000 hours (equivalent Wilma Follette Sequoia (Fresno) . . . . Paul Mitchell to 46.5 full-time employees). BOARD OF DIRECTORS Shasta ...... Ken S. Kilborn CNPS membership is open to all. Sierra Foothills (Tuolumne, Cala- Members receive the quarterly journal, Brett Hall (President); Lauren Brown veras, Mariposa) . . Robert W. Brown Fremontia, the quarterly statewide Bul- (Vice President); Carol Witham (Trea- South Coast (Palos Verdes) ...... letin, and newsletters from their local surer); Laura Camp (Secretary); At David Berman CNPS chapter. Large: Ellen Dean, Arvind Kumar, Brian Tahoe ...... Michael Hogan LeNeve, Nancy Morin, Vince Scheidt, Willis L. Jepson (Solano) ...... Alison Shilling; Chapter Council Mary Frances Kelly Poh Disclaimer: Representatives: Orchid Black, Steve Yerba Buena (San Francisco) . . . . . The views expressed by authors published Hartman Ellen Edelson in this journal do not necessarily reflect established policy or procedure of CNPS, MATERIALS FOR PUBLICATION and their publication in this journal should CNPS members and others are welcome to contribute materials for publication not be interpreted as an organizational in Fremontia. See the inside back cover for submission instructions. endorsement—in part or in whole—of their Staff and board listings are as of April 2012. ideas, statements, or opinions. Printed by Premier Graphics: www.premiergraphics.biz

FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011

ISSUE DATE: OCTOBER 2010 AND JANUARY 2011. PUBLICATION DATE: MAY 2012 CONTENTS USING ALPINE FLORA TO MEASURE ECOLOGICAL EFFECTS OF CLIMATE CHANGE by Jim Bishop ...... 2 CNPS members have helped to establish a network of high elevation monitoring studies to evaluate change in the alpine flora of California that results from climate change.

MANAGING A MOUNTAIN: THE SAN BRUNO MOUNTAIN HABITAT CONSERVATION PLAN by Patrick Kobernus ...... 10 Efficient use of habitat management and monitoring programs will determine how native habitats—and the endangered species they host—respond to current threats facing the site of the nation’s first Habitat Conservation Plan.

RARE PLANTS OF GRIFFITH PARK, LOS ANGELES by Daniel S. Cooper ...... 18 Following a major fire in 2007, biologists have discovered that Los Angeles’ Griffith Park contains a wealth of plant diversity, including numerous rare and locally rare plants.

A PLEA TO PROTECT WALKER RIDGE by Stephen W. Edwards ...... 25 Why Walker Ridge—one of California’s three richest serpentine areas known for its spectacular wildflower displays—deserves protecting.

SERPENTINE SOILS AND WHY THEY LIMIT PLANT SURVIVAL AND GROWTH by Earl B. Alexander ...... 28 Serpentine plants—those on serpentine rocks and in serpentine soils—are unique. Understanding the origins of the rocks and the role of the soils in plant nutrition are keys to understanding the distributions of serpentine plants.

SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA by Hugh D. Safford...... 32 Serpentine soils are uncommon in California but support more than 13% of the state’s endemic plant species. Why is this rare soil type so important to California’s plant diversity?

PLANT GALLS: DESERT TREASURES by Ron Russo ...... 40 The discovery of new plant galls in the Mojave is a reminder of how much we still have to learn about the fabric of desert life.

LEARNING TO READ A LANDSCAPE by Phil Van Soelen...... 46 Becoming a careful observer of nature is a prerequisite to being a successful native plant gardener.

DEVELOPING A SUSTAINABLE MIX FOR SEED GERMINATION USING LOCAL MATERIALS by Jackie Bergquist, Michelle Laskowski, Brianna Schaefer, and Betty Young ...... 50 A report on research by the Golden Gate National Parks Conservancy to develop a seed germination mix as effective as commercial ones, but using only local sustainably produced materials.

CNPS FELLOW: ROGER RAICHE by Phyllis Faber ...... 53

SCOTT FLEMING: 1923–2011 by John Danielson ...... 56

RALPH MILTON INGOLS: 1911–2011 by Leah Price Hawks ...... 57

LETTERS TO THE EDITOR ...... 59

THE COVER: California bristlecone pine woodlands at their upper limit give way to the alpine zone above, a cold, treeless biome being studied carefully for evidence of climate change. Photograph by Stu Weiss.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 1 he early morning is cool and clear, as long shadows of the last and highest-elevation trees stretch Tacross the open slopes ahead. Our team hikes steadily on toward the summit, carrying measuring tapes, compass, level, strings, pin flags and markers, quadrat frames, data binders, cameras, small black- board, and plant press. After pausing on top to distribute the equipment, everyone sets about a task— some lay out the survey system, some assemble . A beautiful world of alpine plants lies at our feet; spectacular views meet our gaze in every direction. It is not yet mid-morning, but we check the sky for the first signs of cloud formation…nothing yet. As soon as the survey areas are defined with colored strings, the work of identifying each plant species and assessing its coverage begins. Meanwhile two people begin the extensive photo documentation protocol.

USING ALPINE FLORA TO MEASURE ECOLOGICAL EFFECTS OF CLIMATE CHANGE by Jim Bishop

ur California team, zone is a sensitive indicator of such lines have risen by over 500 feet. which is working to es- change. Where conditions become less harsh, tablish monitoring sites a greater number of species can sur- in the higher mountains NEED FOR A UNIVERSAL vive. Species richness of many al- Oof the state, is part of an interna- ALPINE-MONITORING pine peaks has increased, and on tional effort to detect and measure PROTOCOL one summit in the European Alps changes in the alpine flora that increased from one to ten species would be expected to occur as the During the past century, scien- over 100 years. That would be ex- climate changes. As an ecosystem at tists have observed changes in al- pected if the cold alpine elevations a climate extreme—one that is very pine flora and in tree line elevation. are warming. But until the last ten temperature dependent—the alpine In Scandinavia, for example, tree years, there had not been a stan-

2 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 dard, universal, and replicable pro- declines steadily with increasing cess for assessing such changes. altitude. Therefore, above tree line In response to the need for bet- elevation, the growing season is sim- ter scientific data and more valid ply too cool to sustain tree growth. comparisons, the Global Observa- But other types of plants persist tion Research Initiative in Alpine in the alpine zone, in spite of cooler Environments (GLORIA, http://www. air temperatures there—plants with gloria.ac.at/) was conceived at the the same temperature-limited basic University of Vienna and established metabolism as trees. How do plants at field sites in Europe in 2001. that live above the tree line do it? GLORIA defines a protocol that can The secret lies in their form and be applied to alpine summits any- stature, allowing them to be warmer where in the world. The alpine zone overall than air temperature would is ideal for detecting biological ef- indicate. There are currently five areas in California fects of global climate changes. Hu- where alpine plant survey work is being The low stature and compact man disturbance there is often mini- conducted as part of the international form of alpine plants reduce cooling mal, it spans nearly all latitudes and GLORIA Project. from the air flow and allow absorp- elevations, and it samples the major tion of sunlight to warm above- climate zones of the world—mari- growing season lengths, latitudes, ground plant parts. Foliage of alpine time and continental, tropical, mid- and precipitation regimes. plants can be sun-warmed above air latitude, and polar. Tree foliage is effectively cooled temperature by some 36o F (so the by the air flow. Tree root zones are temperature of an alpine plant might shaded by the crowns and remain be well over 80o F on a 50o F day). UNDERSTANDING THE close to the average air temperature. The typically sparse distribution of ALPINE ZONE So in effect a tree experiences a cold alpine plants leaves much ground temperature limit dictated by aver- (usually over 50%, often 80% or The story begins with the ques- age air temperature. Average tem- 90%) exposed to solar heating, and tion of why there is an alpine zone. perature in the lower atmosphere their forms transmit more heat to Understanding the environmental stresses on alpine plants and their A team hiking to the Dunderberg target region in the central Sierra . Photograph adaptations is the foundation for by Catie Bishop. All other photographs by the author, except as noted. viewing their responses. The alpine zone is defined as that above the upper tree line—a remarkably con- sistent elevation in a given region, with the trees dropping out com- pletely over elevation increases of just feet. Without the absence of trees there would be no alpine zone. Why should trees meet such a limit to their growth? Answering this question requires understanding how temperature limits plant cell growth, and how it limits tree growth. Plant cell metabolism is very temperature sensitive, slowing to al- most no cell growth at about 42o to 46o F. If the daily mean air tempera- tures (the average for the day) aver- aged over the growing season falls much below about 44o F, trees can no longer subsist. That is a global observation, consistent over differ- ent taxonomic families, elevations,

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 3 Alpine flowers. CLOCKWISE FROM UPPER RIGHT: Cushion Townsend daisy (Townsendia condensata), Sierra penstemon (Penstemon heterodoxus), Pacific (), cushion buckwheat (Eriogonum ovalifolium), White Mountain buckwheat (Eriogonum gracilipes). the ground than do raised crowns. trolled by the distribution of snow- west Research Station, USFS, and Consequently their root system fields. was the location of the first project temperature is above the average of It is expected that at least two sites in the Western Hemisphere. that for shaded sites. In effect, even variables related to climate change Millar remains the force behind all though alpine plants grow at a higher could significantly affect alpine eco- of the California projects and is also elevation than trees, they experience systems—warmer temperatures, and a contributor to other North Ameri- a warmer microclimate. amounts and patterns of precipita- can sites. As beautifully adapted as they tion and snowmelt. Warmer tem- Each GLORIA project is orga- are, alpine plants still live at the peratures can raise the tree line, and nized around a “site” called a target edge of adequate warmth and they will raise the elevations at which a region (TR), and includes a group benefit from the lack of competi- variety of alpine and subalpine plants of three or four summits, spaced in tion from the cold-limited trees, so can be more competitive. Less snow elevation from the highest peaks they are quite vulnerable to tem- or more snow, and changes in pat- down to near tree line. California perature changes. Alpine plant eco- terns of snow deposition and melt GLORIA summits now number 18, systems are also dependent on snow schedule can also alter alpine plant in 5 TRs, from Langley Peak in the distribution. Snow shelters many distributions. southern Sierras to the Carson Range of them from extremes of winter near Lake Tahoe, and at several lo- cold and desiccation. Late-season GLORIA IN CALIFORNIA cations in the central White Moun- snowfields retard growth until they tains. A team of volunteers, students, melt, and upon melting release a GLORIA began in California in and agency employees (who have flush of nutrients and water. Alpine 2004 due to the tireless efforts of been granted release time from their vegetation patterns are strongly con- Connie Millar of the Pacific South- normal duties to help) are conduct-

4 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 ing studies of the alpine ecosystem Enhancements to the basic veys,” observations that cover the and its changes. CNPS volunteers GLORIA protocol have been devel- alpine zone below the GLORIA sum- have been consistent and important oped in California. One of them, the mits and provide a more compre- contributors to the project. “California method,” is now an op- hensive view of the alpine ecosys- Surveys will be repeated at five- tion in the international protocol. tem throughout its entire elevation year intervals to measure change in The observation provides much bet- span. To date, such surveys have the alpine flora. The information that ter quantitative data on plant cover been conducted on four major is gathered will be shared globally, over an area that samples much of slopes, collectively spanning the el- as it is at sites in other countries the variation on the summit. It cov- evation from White Mountain Peak around the world. ers a larger area than the one meter at over 14,000 feet down into the by one meter quadrats, but is not bristlecone-limber pine woodland at subject to the errors of visual esti- 10,800 feet (see protocol details, THE GLORIA SURVEY mates of plant cover on the large page 8). PROTOCOL survey plots. The data gathered from the The White Mountain Research slopes reveals flora that could po- The summit-centered GLORIA Station is now a GLORIA World tentially be moving up toward the protocol outlines a set of survey plots Master Site, where many studies of summits in response to warming. It (successively smaller plots within alpine ecology complement the ba- shows the distribution of each spe- larger ones) that occupy the slopes sic GLORIA surveys. These addi- cies by elevation at a given time. of the four main compass directions tional studies encompass extended Comparing this with the same spe- (N, E, S, and W) on each peak. The plant surveys, tree line and shrub cies’ elevation distribution at a later plots lie in two elevation bands: the line mapping, small-scale tempera- time (for example, ten years later) uppermost band takes in the five ture modeling, insect monitoring, can produce an extremely useful meters (16.4 feet) in elevation just alpine meadow flora, and periglacial measure of just how much the ele- below the peak, and the lower band processes (freeze-thaw cycles). A set vation distribution of that species lies between five and ten meters (33 of weather stations now extends has shifted over time, very possibly feet) below the peak. from the base of the White Moun- rising as the world warms. The materials required to con- tains to the highest summit, provid- duct GLORIA surveys have purposely ing a direct measure of the climate EARLY RESULTS been kept low-tech and therefore are there over a range of elevations— usable worldwide regardless of local important information to relate to What have we learned so far? resources. A compass, tape-measure, any observed ecosystem changes. The initial baseline surveys cannot and a string-level (a builder’s level of themselves reveal change. But they that is attached to a string) are all have provided useful and detailed that are needed for surveying. Col- LOOKING BELOW THE information on the species that are ored string marks the plots, and pin SUMMITS present, their relative abundance, flags have various uses. A one-meter- and their detailed distributions on square quadrat frame with 100 cells A major addition to the summit summits and slopes of the alpine is used for the finest-scale observa- surveys are the “downslope sur- zone in California. tions. All reference points are de- fined by compass direction and dis- Upper edge of bristlecone-limber pine woodland at Patriarch Grove in the White Mountains, tance from the “high summit point.” 1947 and 2005. Dashed line on 1947 photograph indicates road trace corresponding to road Information collected includes in 2005 photograph. Note the many young bristlecone pines within and beyond the woodland in the recent photograph. Most of the global warming in this period took placed after 1970. each species present (including Photographs courtesy of Daniel Pritchett of the White Mountain Research Station. voucher specimens on the first sur- vey); its degree of cover (as a mea- sure of each species’ abundance and its influence on available space, light, air, and water); and the cover of other elements such as rock or soil. The amount of rock and soil indi- cates the potential for expansion of plant cover, and plant cover may change as climate changes (see pro- tocol details, page 6).

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 5 ELEMENTS OF THE GLORIA PROTOCOL he main objective of the GLORIA protocol is to record the plants that grow on the summit, to note roughly T where they are distributed around the peak, and to estimate how much area each species covers. That requires looking at both large-scale and small-scale plots, and on the differently-oriented slopes.The plots are outlined with colored string, and field workers examine each plot carefully for the plants that occur there. A set of survey plots is laid out on the slopes of the four main compass directions or “aspects” (N, E, S, and W). They lie in two elevation bands: the uppermost band spans the five meters in elevation just below the peak, and the lower band lies between five and ten meters below the peak. In each summit area section (SAS)—the area on one aspect and within one elevation band—the plant cover is visually estimated. In the 1m x 1m quadrats (outlined with a frame containing 100 cells), cover is estimated by noting the occurrence of each species in each A diagram of the survey plots as they would appear on the south slope of of the cells. Each group of four quadrats is this summit (HSP is the high summit point). Orange marks the perimeters at the 5-meter and 10-meter levels; pink marks the boundaries of the centered on a main compass direction at the south-slope plots; yellow is the California Method diamond; green is the 5-meter perimeter. grid that locates the one-meter quadrats. Over 60 photographs are taken on each summit, from broader photos of the SAS to the square meter of each quadrat. The photos also document the plants and their coverage, as well as the survey system reference points for placing resurveys. To learn how the actual temperatures and snow cover are changing, a temperature recorder is buried ten centimeters deep on each aspect. Temperature is recorded approximately hourly for several years. The recorder is then dug up, and the records are recovered. The soil-temperature curve has a narrower daily temperature range than the surface-temperature curve, but has a similar average. Changes in average air temperature will be reflected in soil temperature. But when snow covers the ground, the soil temperature remains essentially constant at 32o F, and in that way reveals precisely the periods of snow cover.

BELOW LEFT: Identifying plants. BELOW RIGHT: Searching for inconspicuous plants. Photographs by Scott Sady/TahoeLight.com.

6 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 AN ADDITIONAL PROTOCOL: THE CALIFORNIA METHOD 10 X 10-meter “diamond” is cen- A tered in each aspect on the 5-meter perimeter. Within the plot, 400 sample points set on a half-meter grid produce useful quantitative data on plant coverage at the mid-scale (surveys prior to 2009 used 100 points). Estimating plant cover visually is difficult, especially for plants that have low cover values, and such esti- mates are of very limited accuracy. The quadrat measurements are much more accurate than visual estimates, but they cover only small areas. The 10m X 10m areas yield good accuracy over intermedi- ate areas.

TOP: Taking data on the quadrat grid. • ABOVE LEFT: Documenting photograph points. • LEFT: Leveling the string from which to determine the 5-meter and 10- meter perimeters. Photographs by Catie Bishop. • ABOVE: Measuring the distance to reference points, which are used to guide the placement of resurveys.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 7 In California, four of five target regions have been resurveyed, and two of the downslope surveys in the White Mountains have been re- peated. Many of the re- peat surveys show in- creased species richness. The recent downslope resurveys also showed an increase in the num- ber of annual species (perennial habit is best suited to the demands of the alpine environ- ment, and alpine an- nuals are uncommon). Outline of one segment, 1 meter wide and 10 meters long, centered on the white tape, of the 100-meter belt transect at that elevation. All species are noted within However, that may be due, in part, the segment. Sample point positions for plant species or cover type are indicated to a wetter-than-normal year. on this photograph with red circles. “Frequency” refers to the total number of Findings over the ten years from segments a species occurs in; “Cover” refers to the number of sample points that 1994–2004 from the European Alps contact a given species or cover type. have shown reductions in cover of the highest elevation species (those DOWNSLOPE SURVEY PROTOCOL in the zone of permanent snow, or the “nival” zone), with lesser reduc- ownslope surveys are conducted on some of the slopes below tions and some increases within the the GLORIA summits, into the subalpine woodland. Meter- D alpine subzones below that. These wide belt transects along the elevation contour are spaced every 25 changes show that the highest plants meters (82 feet) in elevation. Each transect is 100 meters long (100 are losing ground to plants from square meters total area), divided into 10-meter segments. All spe- lower elevations, at least over a re- cies present in each 10-meter segment are recorded. At cent decade. sample points spaced every half meter (400 for the Even if the changes observed in entire transect) the plant species or other cover type is the GLORIA resurveys are signifi- identified. The downslope transects duplicate, in area cant, it is not possible to attribute and sampling density, the 10m X 10m California- change over five years to long-term method plots on the summits. climate trends. Inter-seasonal varia- tion can easily affect the flora rep- LEFT: A pointer that marks two sample points, one at each end, is moved along and placed every one-half meter along the tape. • resented in a given year. The value BELOW: Botanists look closely for small plants, since it is easy to of the current comparisons is that miss some of the Alpine species, which can be quite tiny. they demonstrate the potential of

8 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 the protocols to reveal change. risen 500 feet, and several common REFERENCES However, only changes in a direc- alpine plants have been reduced in tion consistent with a warming cover wherever the Artemisia has Koerner, C. 2003. Alpine Plant Life, world, sustained over two or more encroached. Aerial photo compari- Functional Ecology of High Mountain Ecosystems, 2d. ed. Springer-Verlag, decades, will indeed be compelling sons from the late 1940s into the Berlin, Heidelberg, New York. evidence. 2000s show bristlecone pine wood- IPCC Working Group 1. 2007. Climate Current data is also being com- land filling in along its upper mar- Change 2007, the Physical Science pared to earlier botanical data from gins and young trees spreading Basis. Cambridge University Press, the White Mountains. For example, upslope. Cambridge, MA. the uppermost occurrence on gra- Much remains to be learned, and White Mountain Research Station, nitic substrates of dwarf sagebrush only time will reveal long-term GLORIA website: http://www.wmrs. (Artemisia arbuscula) as reported in change. But the baseline data being edu/projects/gloria%20project/ 1961 has been compared to the re- established, and the early look at default.htm cent upper shrub line surveys and changes, are an important contribu- downslope surveys. Based on those tion to assessing the impacts of cli- Jim Bishop, 1144 Mount Ida Road, observations, which span a 50-year mate change on the alpine ecosys- Oroville, CA 95966, cjbishop1991@ interval, this species’ upper limit has tem across the world. sbcglobal.net

he team is finishing up its work on the summit. Strings are wrapped on cardboard holders, data T books carefully packed away. We enjoy last views of the magnificent landscape, and begin the hike down. Beginning with a few morning cumulus over the highest peaks, the clouds have now become tall and massive, bright against the mountain sky. A faint roll of thunder is heard from just a few miles away. It is good to be on our way down. Back at camp we’ll finish up some challenging plant identifications and file the data sheets. We’ll gather the equipment for the next day’s work, and vow to begin early enough to finish before storms end the field day.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 9 MANAGING A MOUNTAIN: THE SAN BRUNO MOUNTAIN HABITAT CONSERVATION PLAN by Patrick Kobernus

n 1982, the first Habitat Conserva- conserved habitat.1 As of 2012, most panding populations of invasive spe- tion Plan (HCP) in the nation was of the development on the moun- cies, continue to overtake grassland approved for San Bruno Moun- tain has been completed. habitat on the mountain. The future tain. Over the past 30 years most Management of invasive species conservation of the endangered spe- Iof the mountain, approximately to protect the endangered species cies and their habitats, now more 2,830 acres (82%), has been con- habitat on the mountain has been than ever, depends upon implemen- served as habitat for three endan- largely successful over the past 30 tation of a more comprehensive habi- gered butterfly species, wildlife, and years. However, coastal scrub suc- tat management program to protect plants; and approximately 350 acres cession, in combination with ex- these species for future generations. (10%) of the mountain has been de- veloped. An additional 260 acres (8%) of land remains undeveloped, 1 Most of the remaining “unplanned parcels” will likely be conserved due to these lands and it is likely that a majority of this being located on very steep slopes with no infrastructure (roads, utilities) for development. land will ultimately be set aside as The HCP requires that a minimum of 40% of this land be dedicated as conserved habitat.

MOUNTAIN FLORA AND FAUNA

San Bruno Mountain is located in northern San Mateo County, ad- jacent to San Francisco. It consists of approxi- mately 2,830 acres of open space, and is bor- dered by the urban and suburban portions of Daly City, South San Francisco, Colma and Brisbane. Though it is isolated by urbanization, the mountain is consid- ered the northernmost part of the Santa Cruz Mountains. The famed botanist James Roof asserted that San Bruno Mountain supports one of the last and the most expansive areas of a unique and highly diverse grassland and shrubland flora, The San Bruno Mountain Habitat Conservation Plan—adopted in 1982 and the first HCP in the nation— which he referred to as is responsible for protecting host plants for three endangered butterflies, and conserving the area’s “Franciscan” (Edwards diverse native flora and fauna. All photographs by the author unless otherwise indicated. 2000). This Franciscan

10 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 flora was once common through- rocky outcrops, roadcuts, and on out the hills of San Francisco but cut slopes. has been almost entirely destroyed The mountain is not only in the city by development and home to three endangered butter- planting of nonnative trees. fly species but also supports a wide The grassland on San Bruno diversity of other native flora and Mountain is actually a combina- fauna. Many varieties of wildflow- tion of different types of grass- ers can be found on the moun- lands intergrading and sharing tain, including coast rock cress some of the same wildflower and (Arabis blepharophylla), Pacific shrub associates. Grassland types stonecrop, varied-color lupine, vary on the mountain de- Johnny jump up, gold- pending on elevation, ex- fields (Lasthenia cali- posure, and soil type. The fornica), shooting stars dryer southern exposures (Dodecatheon hendersonii), tend to have stands of blue larkspur (Delphin- purple needle grass (Nas- ium decorum), farewell to sella pulchra), while the spring (Clarkia rubicun- more fog-shrouded grass- da), and owl’s clover (Cas- lands near the summit tilleja densiflora), among are dominated by Califor- many others. Each patch nia fescue (Festuca califor- of grassland is a uniquely nica), red fescue (F. rubra), beautiful “natural garden” and fescue (F. ida- that has been constructed hoensis). There are also through the forces of na- stands of California oat ture and time. Each March grass (Danthonia califor- through June, the grass- nica), blue wild rye (Ely- lands and wildflowers mus glaucus var. glaucus), emerge and change into Pacific reed grass (Calamagrostis new combinations of color and nutkaensis), June grass (Koeleria beauty as the season progresses. macrantha), and tufted hair grass There are several rare plant (Deschampsia cespitosa ssp. holci- species including two that are formis). state and/or federally listed, San As a biologist hired to moni- Bruno Mountain manzanita (Arc- tor the three endangered butter- tostaphylos imbricata) and San fly species on San Bruno Moun- Francisco lessingia (Lessingia tain for 13 years, I can attest to its germanorum). There are also sev- unique beauty. Each spring and eral California Rare Plant Rank summer, I would hike the moun- 1B species (formerly CNPS List tain repeatedly while I recorded 1B) such as Montara manzanita my observations of the mission blue, TOP: A freshly emerged male mission blue (Arctostaphylos montaraensis), Paci- butterfly (Icaricia icarioides missionensis), San Bruno elfin, and callippe found in San Bruno Mountain’s grassland fic manzanita (Arctostaphylos pa- silverspot butterflies, and the status habitat. It is an endangered species. • cifica), Diablo helianthella (Helian- of their grassland habitats. MIDDLE: A mission blue larva feeding on thella castanea), San Francisco spine- The San Bruno elfin’s host plant, silver lupine (Lupinus albifrons var. collinus) flower (Chorizanthe cuspidata var. Pacific stonecrop (Sedum spathuli- on the west peak of San Bruno Mountain. cuspidata), and San Francisco cam- • BOTTOM: A San Bruno elfin larva foraging folium), grows in coastal prairie and on the flowerheads of its host plant, Pacific pion (Silene verecunda ssp. verecun- on rocky outcrops and roadcuts. stonecrop (Sedum spathulifolium) near the da). Other rarities include an arach- The Callippe silverspot’s host plant, summit of San Bruno Mountain. nid, incredible harvestman (Bank- Johnny jump up (Viola pedunculata), sula incredula); a solitary bee (Du- grows in coastal prairie and in non- ied-colored lupine (L. variicolor), fourea stagei); and several range- native annual grasslands. The mis- and summer lupine (L. formosus var. limited endemic plants. sion blue’s host plants, silver lupine formosus) grow within coastal prai- Plant communities on the moun- (Lupinus albifrons var. collinus), var- rie, nonnative annual grassland, tain include northern coastal scrub,

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 11 coast live oak woodland, coastal ter- This mechanism has been used Bruno elfin butterfly’s habitat has race prairie, freshwater marshes and as a tool for settling land disputes been protected. seeps, central coast riparian scrub, by allowing landowners to econo- The development permitted nonnative gorse and broom scru- mically develop portions of their through the HCP is primarily rele- blands, nonnative eucalyptus for- properties, while simultaneously en- gated to the lower slopes of the est, and nonnative annual grass- suring long-term protection of en- mountain, thereby protecting the land. The most dominant vegeta- dangered species through dedication majority of higher quality butterfly tion on the mountain is northern of conservation areas, habitat man- habitat on the upper ridges and hill- coastal scrub and nonnative annual agement and monitoring, and/or tops. For the callippe silverspot, this grassland. other mechanisms. As of 2010, over was critical, because this species re- 700 HCPs have been permitted in quires hilltops for mate selection. the U.S. by the U.S. Fish and Wild- Male callippes stake out territories FIRST HABITAT life Service (USFWS 2010). on the highest hilltops or ridgelines CONSERVATION PLAN The primary purpose of the San available to encounter and attract IN U.S. Bruno Mountain HCP is to protect females for mating. the grassland habitat that supports The conservation areas also in- Since 1982 the mountain has the three endangered butterfly spe- cluded protection of several rare been the site of the first Habitat cies, while allowing limited devel- plant species, with the exception of Conservation Plan (HCP) in the opment to occur. Prior to the for- most of the San Francisco lessingia nation. HCPs were created as a mation of the HCP, approximately and San Francisco spineflower pop- mechanism to balance private prop- 1,950 acres of land on San Bruno ulations, which are located on pri- erty rights and endangered species Mountain had been purchased and/ vate property on the west side of protection, by allowing limited “tak- or donated to create the San Bruno the mountain. These populations are Mountain State and still extant, however development County Park. This and invasive species are potential land contained vir- threats. tually the entire The HCP specified the impor- habitat for the San tance of management and monitor- Bruno elfin butter- ing of the butterflies’ habitat and fly on the mountain. provided a funding mechanism to However, it did not support these activities—an annual include the prime monetary assessment on every resi- habitat areas for the dence and business that is built mission blue and within the HCP boundary. This as- callippe silverspot sessment rises with the annual in- butterflies, which flation rate, and has provided a con- were located on the sistent level of funding since the cre- eastern portions of ation of the HCP. the mountain. The HCP fund is managed by The HCP pro- the HCP Trustees (the City Manag- vided a mechanism ers of Daly City, Brisbane, and South View of housing development and mission blue butterfly host by which an addi- San Francisco, and the San Mateo plant, silver lupine (Lupinus albifrons var. collinus) on the tional 800+ acres County Manager). Monitoring and southeast ridge of San Bruno Mountain. Development is no longer the most serious threat to the grassland habitat on the of habitat would habitat management is implemented mountain. A greater threat is coastal scrub succession and the be conserved and by San Mateo County Parks Depart- expansion of invasive species. added to the Park, ment, and implementation of the and all of the con- plan is overseen by the U.S. Fish ing” (destruction) of endangered served land within the park would and Wildlife Service. species and their habitat, provided be managed and monitored for the that the taking will not appreciably endangered species as well as for reduce the likelihood of the sur- other native flora and fauna. Within THREATS FROM INVASIVE vival and recovery of the species in the current 2,830-acre conservation SPECIES the wild, as specified in section area, approximately 90% of the mis- 10(a)(1)(B) of the federal Endan- sion blue and callippe silverspot but- In 1982, an assortment of ag- gered Species Act. terflies’ habitat, and 100% of the San gressive invasive plant species were

12 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 identified and mapped on the moun- implementation of ongoing habitat proven to be successful in maintain- tain. Gorse (Ulex europaeus), the management with an emphasis on ing most of the habitat for the en- most aggressive of these plants, was controlling invasive species became dangered species over the 30-year introduced to the mountain in the an important component of the San span of the HCP. While the reduc- 1930s and by 1982 had expanded to Bruno Mountain HCP. Due to the tions of large infestations has been cover several hundred acres of the lack of information on the feasibil- effective, it is the less noticeable habi- mountain. Based on the rate of ex- ity of habitat restoration at the time tat maintenance work that is done pansion of this plant and that of of the inception of the HCP, the year in and year out by HCP work other invasive weeds, combined with HCP’s primary goal has been focused crews and volunteer groups that illegal off-road vehicle use and on the conservation and manage- serves to protect the majority of the coastal scrub succession, it was esti- ment of existing habitat for the but- butterfly habitat on the mountain. mated that the habitat for the en- terflies (San Bruno Mountain HCP dangered butterfly species on the Steering Committee 1982). HABITAT RESTORATION mountain could be completely wiped Though the HCP has often been out within 50 to 200 years (San criticized for the lack of restoration Restoration of mission blue and Bruno Mountain HCP Steering Com- work that has been conducted, the callippe silverspot habitat is re- mittee 1982). strategy of focusing efforts on pro- quired by the HCP within areas that As a result, the funding and tecting the existing habitat has were disturbed by grading activi-

Extent of grasslands on San Bruno Mountain as mapped in 2004. Map produced by TRA Environmental Sciences and provided courtesy of San Mateo County Parks Department.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 13 14 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 ties adjacent to the developments. Mateo County Parks Department grasslands on San Bruno Mountain These areas have steep slopes that 2008). This corresponds to a loss of was evaluated in 2002 using live were engineered to protect against over five acres of grassland per year. and dead above-ground biomass landslides. Restoration of butterfly The expansion of coastal scrub veg- measurements. Values measured habitat has not been successful on etation and corresponding loss in within the grasslands on the south most of these slopes, due to the grassland has been documented in slope of the mountain prior to difficulty in establishing native many regions of California (Murray experimental grazing treatments, plants in poor soil conditions. This 2003, McBride and Heady 1968), showed live and dead above ground is especially true for the host and and is often the result of the re- biomass levels of 5,000 to 9,000 nectar plants for the callippe moval of grazing and/or burning lbs/acre. A large proportion of this silverspot butterfly. Propagating and from a grassland ecosystem. was from thatch. As a comparison, replanting of the callippes’ host Historically cattle grazing and the recommended ranges for Re- plant Johnny jump up has shown brush burning by local ranchers re- sidual Dry Matter (live biomass) in to be expensive, with very little suc- sulted in the control of coastal scrub coastal prairie grasslands with mini- cess to date. Johnny jump up is vegetation on San Bruno Mountain, mal woody plant cover ranges from difficult to grow under nursery con- but also facilitated the spread of in- 1,200 to 2,100 lbs/acre (UC Davis ditions, and has had a very low vasive plant species. Invasive grasses 2002). survival rate when transplanted into such as ripgut brome (Bromus dian- The reduction in wildfires, re- restoration sites. For the mission drus), velvetgrass (Holcus lanatus), moval of grazing animals in the early blue however, its host plants (espe- and invasive herbaceous weeds such 1960s from the mountain, and at- cially silver lupine) have been re- as fennel (Foeniculum vulgare), wild mospheric nitrogen pollution are all established on several restoration radish (Raphanus sativus), and oxa- likely factors contributing to the pro- slopes because these plants are lis (Oxalis pes-caprae) have prolifer- liferation of invasive plants, build- adapted to disturbed rocky slopes ated because of the ability of these up of thatch, and brush succession with thin or poor soil conditions. species to rapidly expand into grass- on the mountain. lands (San Mateo County Parks Department 2008). Atmospheric FUTURE MANAGEMENT CURRENT THREATS AND sources of nitrogen pollution (smog) MANAGEMENT may also be contributing to the The San Bruno Mountain HCP spread of these invasive grasses and has been an experiment—the first While efforts have been success- weeds within the grasslands (Weiss of its kind—to protect endangered ful in reducing the large, woody in- 2006). species habitat while allowing lim- vasive species on San Bruno Moun- As more and more weeds prolif- ited development. For 30 years the tain, control work has been less ef- erate and die back, the resultant plan has been a qualified success in fective at stemming the tide of coastal accumulation of live and dead bio- that all three of the endangered but- scrub succession. An independent mass (thatch) reduces the amount terfly species on the mountain con- analysis of almost 20 years of but- of light reaching the soil surface, tinue to be locally abundant. How- terfly data collected over the course suppressing the growth of native ever, management of the conserva- of the HCP revealed that the overall grassland plants. Increased mois- tion areas will need to adapt to distribution of the mission blue and ture retention from the shade cre- changing conditions and address callippe silverspot butterflies re- ated by thatch may also facilitate problems such as coastal scrub suc- mained stable. However, geographic the expansion of coastal scrub into cession and invasive weeds in a more areas of concern were identified for the grassland areas. Furthermore, comprehensive way. The 2008 San each species. where invasive control work has Bruno Mountain Habitat Manage- For the period between 1982 and been done for decades, there is a ment Plan spells out in detail the 2004, San Bruno Mountain lost an significant build-up of thatch from priority areas to protect, the current estimated 122 acres (8.6%) of grass- old stalks of fennel, broom, and and emerging threats to the moun- land habitat. This was primarily due gorse plants that were left to decay tain’s habitats, and the methods for to coastal scrub succession within in place. monitoring and management to ad- the HCP conservation area (San The level of thatch within the dress these threats.

FACING PAGE, TOP: Buckeye Canyon in 1986. This photograph was taken approximately 25 years after cessation of cattle grazing on San Bruno Mountain. • BOTTOM: Buckeye Canyon in 2006. This photograph was taken approximately 45 years after the cessation of grazing. Coastal scrub vegetation is overtaking grasslands at a rate of approximately five acres per year. Photographs courtesy of TRA Environmental Sciences and San Mateo County Parks Department.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 15 View of the ridge above Owl Canyon near the summit of San Bruno Mountain. The area contains prime grassland habitat, including native wildflowers and shrubs, for the Callippe silverspot and San Bruno elfin butterflies, both endangered.

Until 2010, threats to native pected to occur from global climate Bruno Mountain that supported the habitats on San Bruno Mountain change, it is important to preserve alternative host plant summer lu- could not be addressed comprehen- as much potential grassland habitat pine were unaffected by the fungus, sively given the existing manage- as possible to buffer the endangered and mission blue butterfly numbers ment budget. However, as a result species from occasional large-scale in these areas remained stable. In of an agreement reached between declines in habitat quality. For ex- the subsequent 14 years, silver lu- developers, the city of Brisbane, San ample, in the extremely wet El Niño pine plants have rebounded signifi- Mateo County Parks Department, year of 1998, the numbers of mis- cantly, as have the mission blue and the U.S. Fish and Wildlife Ser- sion blue butterflies on San Bruno numbers in the areas impacted by vice, an additional four million dol- Mountain declined markedly in ar- the fungus. Protecting areas of dif- lars will be generated through de- eas where silver lupine was the domi- ferent habitat quality, slope aspect, velopment fees and placed into an nant host plant. Silver lupine expe- and within different microclimates endowment for the mountain. Once rienced a widespread die-off due to is important because habitat quality collected, these funds would increase a fungal infestation brought on by can be expected to fluctuate over the HCP annual budget for habitat the excessive wet soil conditions. time, due to plant senescence and management by two to three times This impact was observed through- climatic factors, and these fluctua- its former level. These funds need to out the range of the species (includ- tions may become more extreme in be used to manage more grassland ing the Marin headlands and at Twin the future. areas on the mountain. Peaks, San Francisco). While prescribed burning may With accelerated changes ex- In contrast, habitat areas on San continue to be difficult to imple-

16 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 ment on San Bruno Mountain due to mental groups has been on fighting Journal of the Regional Parks Botanic public safety concerns, grazing and/ development on San Bruno Moun- Garden, Volume 11, Number 2. Oc- or mowing should be implemented tain, under the assumption that de- tober 2000. Published by East Bay to reduce vegetation fuel loads be- velopment is the major threat to Regional Park District, Oakland, CA. tween parkland areas and homes and the endangered species. In reality, McBride, J., and H.F. Heady, 1968. In- businesses. Stewardship grazing or though, the permitted development vasion of Grasslands by Baccharis Pilularis. Journal of Range Manage- mowing of 100–500-foot buffer has impacted approximately 10% ment. 21:106-108. zones on regular intervals would re- of the mountain and was relegated Murray, M.P., 2003. Losing ground: wil- duce fuel loads near populated areas to lower slopes, generally of lesser derness meadows and tree invasion and could potentially allow for the habitat value. The development is over a 55-year period in California’s safe use of controlled burns in some now nearing completion. The only Klamath Range. Pp. 81-97, In: K.L. areas of the mountain. Also, grazing way to protect the endangered spe- Mergenthaler, J.E. Williams, and or mowing within lower elevation cies and the plant communities of E.S. Jules (eds). Proceedings of the areas between parklands and devel- San Bruno Mountain for future gen- Second Conference on Klamath- opments would not impact the more erations will be to manage the re- Siskiyou Ecology. Siskiyou Field intact stands of coastal prairie, which maining conserved habitat more Institute, Cave Junction, OR. are more concentrated on the upper effectively. San Bruno Mountain Habitat Conser- slopes of the mountain. The 2008 Habitat Management vation Plan Steering Committee, 1982. San Bruno Mountain Habitat Grazing, mowing, and/or burn- Plan for San Bruno Mountain estab- Conservation Plan, Volumes I and II. ing will need to be applied to ad- lished a goal of maintaining between Chaired by the County of San Mateo. dress scrub succession and invasive 1,200–1,800 acres of native and non- Plan prepared by Thomas Reid As- weed infestations on San Bruno native grassland on the mountain. sociates, November. Mountain. These tools will need to Currently the area of grassland is San Mateo County Parks Department, be used in combination with other approximately 1,250 acres, but it is 2008. San Bruno Mountain Habitat weed control methods to manage decreasing at a rate of approximately Management Plan, 2007. Prepared areas effectively. There must be a five acres per year. Slowing the rate in Support of the San Bruno Moun- prescriptive approach that is tailored of coastal scrub succession and in- tain Habitat Conservation Plan. Re- in timing, duration, and frequency creasing the amount of grassland will vised 2008. San Mateo County Parks to each area of the mountain, de- require that brush control programs and Recreation Division. March pending upon the grassland type, be implemented sooner rather than 2008. University California at Davis, 2002. surrounding terrain, presence of rare later. California Guidelines for Residual and endangered species, and public The San Bruno Mountain story Dry Matter (RDM) Management on safety concerns. is not unique: brush succession and Coastal and Foothill Annual Range- invasive species have been negatively lands. Rangeland Monitoring Series, CONCLUSIONS impacting grasslands and meadows Publication 8092. University of Cali- throughout California for several fornia Davis, Division of Agriculture Over the years, the San Bruno decades. The San Bruno Mountain and Natural Resources, California Mountain HCP has received a sub- HCP is unusual, however, in that it Rangelands Research and Informa- stantial amount of criticism from has had a mandate and funding to tion Center. environmental groups regarding the address these issues for 30 years. It U.S. Fish and Wildlife Service, 2010. lack of successful habitat restora- will take a coordinated effort on the U.S. Fish and Wildlife Service, Con- tion on the mountain. In contrast, part of biological monitors, habitat servation Plans and Agreements Da- tabase. Website accessed October 10, resource managers have emphasized managers, oversight agencies, envi- 2010. http://ecos.fws.gov/conserv_ the positive aspects of the HCP and ronmental groups, and the commu- plans/public.jsp how it has worked to protect the nity to work cooperatively and cre- Weiss, S.B., 2006. Impacts of nitrogen endangered species habitat and na- atively to ensure that the mountain’s deposition on California ecosystems tive plant communities. The reality endangered species and unique and biodiversity, California Energy is that both groups are right. The Franciscan flora are protected for Commission Report. Website publi- restoration work has been largely the next 30 years. cation accessed on June 25, 2007. unsuccessful on the graded slopes, http://www.creeksidescience.com/ while the habitat management has publications.html REFERENCES been successful in protecting the endangered species populations Edwards, S.W., 2000. The Franciscan Patrick Kobernus, 1072 Geneva Avenue, within the conservation areas. Region. Transcript of a Lecture by James San Francisco, CA 94112, crecology@ The primary focus of environ- Roof ca. 1975. The Four Seasons, gmail.com

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 17 View southeast from block F8 of Griffith Park (see Figure 2) showing downtown Los Angeles (at right) and the Los Angeles River (center). RARE PLANTS OF GRIFFITH PARK, LOS ANGELES by Daniel S. Cooper

ising more than a thousand feet ged interior is still surprisingly wild, such landmarks as the Griffith Ob- off the floor of the Los Ange- owing to both its steep topography servatory and the Los Angeles Zoo, les Basin, Griffith Park divides and lack of established trails. none of which sustained damage. the eastern San Fernando Val- Just a few weeks earlier, on March Rley from the coastal plain of Los 30, an arson fire had swept up to- Angeles and protects more than WILDFIRE PROVIDES ward the northwestern corner of the 4,000 acres of rugged slopes and IMPETUS FOR NEW park, burning a large area of high- canyons at the eastern end of the RESEARCH elevation atop Cahuenga Santa Monica Mountains. Although Peak, the highest point in the east- portions of the park are heavily used, On May 8, 2007, an arson- ern Santa Monica Mountains. While with golf courses, grassy picnic ar- caused wildfire burned about 20% these fires radically altered portions eas, and other attractions situated of the park, essentially the entire of the park’s vegetation—at least mainly at the park’s borders, its rug- southeastern corner, threatening temporarily—their more lasting

18 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 contribution may have been the in- because of its rugged topography Mountains. Included in this list were creased awareness they elicited from and the fact that most readily acces- both CNPS rare taxa (California local conservationists and city staff sible hiking areas (mainly fire roads) Native Plant Society 2010), as well alike, most of whom had never ex- have been degraded by weed abate- as a group of species we felt were plored the park. Suddenly, the park ment practices and non-native veg- significant, either because they are seemed like a worthy subject of im- etation. rare in the region, or because if mediate ecological exploration, per- found, they would represent dis- haps because it was almost com- SURVEY LAUNCHED junct occurrences from populations pletely reduced to ash-covered farther west in the Santa Monica slopes. To remedy this information gap Mountains, or to the north in the Just prior to the fires and con- and to identify critical areas for plant Verdugo and/or San Gabriel Moun- tinuing to the present, a collection conservation, an informal, volun- tains. of neighborhood groups and the Los teer-based “Griffith Park Rare Plant I also searched for taxa known Angeles/Santa Monica Mountains Survey” was launched in spring 2010 in the area only from historical col- Chapter of CNPS funded several (Cooper 2010). The botanical his- lections (Consortium of California projects aimed at gathering baseline tory of an area can be a challenge to Herbaria 2011), though some of biological data on its plants and ani- uncover, even for trained research- these were likely collected in areas mals. These were carried out in con- ers. Most parts of California remain no longer undeveloped (e.g., the Los junction with the development of a very poorly known botanically, in- Angeles River and “Providence wildlife management plan funded cluding sites within urban areas. Of- Ranch,” since converted to Forest by the City of Los Angeles (Cooper ten there are a few local experts who Lawn Memorial Park). A small group and Mathewson 2009). While the have botanized in an area flora of the Santa Monica Moun- and have kept at least Figure 1. Locations of Rare Plants in tains as a whole has been well docu- some field notes (or to- Griffith Park mented (Wishner 1997; Gibson and day, digital photographs), Prigge 2003; California Native Plant and can help put together Society, undated), that of the far a list of species likely to eastern end of the range is compara- occur based on their ex- tively less well-known than the large, perience in the region. protected areas to the west such as Such observations may Topanga Canyon State Park and the be supplemented by plant Santa Monica Mountains National collections in local her- Recreation Area. baria, which are often the Aside from scattered references only record of species (e.g., McAuley 1985), the flora of long gone from urban Griffith Park had never been syn- sites such as native wild- thesized, although dozens of speci- flowers. mens had been collected here since Through my own the mid-1800s, mostly prior to 1950 fieldwork conducting (Consortium of California Herbaria wildlife surveys in the 2011). The one available checklist park since 2007, I was of the plants of the park (Brusha able to identify key habi- KEY: Green: Nevin’s barberry (presumed planted) 2003) does not differentiate between tat features most likely Magenta: Plummer’s mariposa lily (generalized planted and naturally occurring to hold rare or interest- locations; plant is very widespread) taxa, and presents just a fraction of ing plants, such as rock White: Catalina mariposa lily the true species diversity. Currently outcrops and remote Yellow: Slender mariposa lily an as-yet unpublished checklist of canyons. To begin the Dark Blue: Clay bindweed Light Orange: Humboldt lily the park’s flora stands at more than task of locating rare and Light Blue: Hubby’s phacelia 500 species vouchered or photo- significant species and Red: San Gabriel Mountains leather oak (pending graphed (Cooper, forthcoming). populations, a list of tar- further identification) This is nearly half the known flora get species was devel- of the Santa Monica Mountains and oped with the help of Richard of volunteers used this species list, includes 350 natives. Still, the park Fisher, Bart O’Brien, and Carl along with a map of the park has apparently never been a popu- Wishner, local botanists familiar (Cartifact, Inc. 2007) that had been lar place for botanizing, probably with the flora of the Santa Monica divided into 40 survey blocks, and

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 19 Figure 2. Distribution of Rare and Locally Rare Plant Species in in the Los Angeles area if they per- Griffith Park, by Survey Block sist at all. Three species of mariposa lily occur in the park, and exhibit an interesting, non-overlapping pattern of occurrence, with Plummer’s mari- posa lily ( plummerae) by far the most common. Hundreds of these spectacular pink and yellow lilies were encountered on thin, grav- elly soil along ridges, often in asso- ciation with yucca (Hesperoyucca whipplei), giant stipa (Achnatherum coronatum), coast buckwheat (Erio- gonum fasciculatum), and chamise (Adenostoma fasciculatum). They were often found along small trails and footpaths (including equestrian routes), where this activity appeared to limit non-native grasses, among other plants, that might otherwise overtake the lilies. Catalina mariposa lily was found in pockets of grassland on moist, heavy clay soil near the south-cen- Outline of park is drawn in gray. This grid system follows that used by Cartifact, Inc. tral area of the park, with two large (2007) for a Griffith Park map, and notable natural and physical features are listed within each block. The colors correspond to the number of rare species found within each block, populations discovered that exceed as indicated by the key on the right. For example, pale pink blocks had two to three rare 100 plants. Finally, the slender mari- species, while the darkest red blocks had nine or more. posa lily (Calochortus clavatus var. gracilis) was discovered prior to the sent me photos and notes from their historical specimens (Table 1). (A start of the survey, in a single patch forays throughout spring 2010. description of the California Rare of fewer than 10 individual plants in Plant Ranks, formerly the CNPS a grassy opening in chaparral on a RARE SPECIES FINDINGS Lists, is available at http://www. remote peak near the center of the cnps.org/cnps/rareplants/ranking. park. No other populations of this From this effort, 13 rare species php.) Of the four species believed stunning yellow lily were encoun- (California Rare Plant Ranks 1, 2, extirpated from Griffith Park, only tered, despite much searching. and 4) were identified as occurring one, the Brewer’s redmaids (Calan- Humboldt lily (Lilium humboldtii or having occurred in or adjacent to drinia breweri), has been collected var. ocellatum) appears to be con- Griffith Park. Of these, nine were recently in the region near the park fined to just four oak-shaded can- found to be extant (still in exist- (Verdugo Mountains); the remain- yons in the park, with most plants ence), and four are known only from ing species are apparently very rare (50+ individuals) located in Brush

LEFT: Plummer’s mariposa lily is found frequently on steep ridges on eroding soil throughout Griffith Park, Los Angeles. All photographs by the author unless otherwise specified. • MIDDLE: Chaparral pea emerging atop Cahuenga Peak after 2007 fire in Griffith Park. • RIGHT: The rare Humboldt lily is found beneath oaks in several deep canyons in Griffith Park. Photograph by Gerry Hans.

20 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 TABLE 1. SPECIAL STATUS PLANTS AT GRIFFITH PARK

Representative Park status Species Legal status specimen1 (40 survey blocks)

Nevin’s barberry Federal: SBBG37272 Probably introduced; Berberis nevinii Endangered found in 5 survey State: blocks Endangered CA Rare Plant Rank 1B.1 Brewer’s redmaids CA Rare Plant JEPS17234 Historical Calandrinia breweri Rank 4.2 specimen(s); no modern record Catalina mariposa lily CA Rare Plant RSA15196 8 survey blocks; Calochortus catalinae Rank 4.2 (“Cahuenga Pass”) heavy clay soil slender mariposa lily CA Rare Plant (photograph 1 survey block, Calochortus clavatus Rank 1B.2 only) < 10 plants var. gracilis Plummer’s mariposa lily CA Rare Plant LA29033 13 survey blocks, Calochortus plummerae Rank 1B.2 gravelly ridges Clay bindweed CA Rare Plant UCR216375 1 survey block, Convolvulus simulans Rank 1B.2 < 10 plants many-stemmed liveforever CA Rare Plant RSA397814 Historical Dudleya multicaulis Rank 1B.2 specimen(s); no modern record large-leaved filaree CA Rare Plant RSA390952 Historical Erodium macrophyllum Rank 4.2 (“Providencia specimen(s); no Ranch”) modern record southern California CA Rare Plant JEPS58691 Common park-wide; black walnut Rank 4.2 not mapped Juglans californica Humboldt lily CA Rare Plant (photograph 6 survey blocks, Lilium humboldtii Rank 4.2 only) mesic canyons var. ocellatum Hubby’s phacelia CA Rare Plant UCR209589 2 survey blocks, Phacelia cicutaria var. hubbyi Rank 4.2 sedimentary rock Cooper’s rein-orchid CA Rare Plant RSA382793 Historical Piperia cooperi Rank 4.2 (“Providencia Ranch”) specimen(s); no modern record San Gabriel Mountains CA Rare Plant RSA652868 1 survey block leather oak Rank 4.2 Quercus durata var. gabrielensis

1 Location is “Griffith Park” unless otherwise indicated. RSA = Rancho Santa Ana Botanic Garden Herbarium SBBG = Santa Barbara Botanic Garden LA = UCLA Herbarium JEPS = Jepson Herbarium, UC Berkeley UCR = UC Riverside Herbarium

Sources: Consortium of California Herbaria 2011; Cooper 2010.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 21 Another difficult-to-assess rare flora, and more probably await dis- plant in the park is the San Gabriel covery. The only known popula- Mountains leather oak (Quercus tion of dwarf brodiaea (Brodiaea durata var. gabrielensis, California terrestris var. kernensis) in the east- Rare Plant Rank 4.2), which is ern Santa Monica Mountains (A. known from a single collection in Gibson, pers. comm.) was also dis- 1991. However, more study and col- covered by a volunteer during the lection of various scrub oaks in the survey. park are needed, particularly since Uncommon species reaching a Griffith Park may lie within an in- local distributional limit near the trogression zone between the flora park include erect goldenaster (He- Dwarf broadiaea, a locally rare plant other- wise unknown in the eastern Santa Monica of the Santa Monica and San Gabriel terotheca sessiliflora var. fastigiata), Mountains, was discovered just outside the Mountains. This variety appears to the latifolium subspecies of the wide- park boundary in 2010. be otherwise unknown in the Santa spread California fuchsia (Epilobium Monica Mountains. canum), and valley cholla (Cylindro- Canyon in the southwest corner. A map showing the distribution puntia californica var. parkeri). A few Hubby’s phacelia (Phacelia cicutaria of the above rare species in the park common wildflowers in the park var. hubbyi) was found on exposed, is provided in Figure 2 on page 20. show more of a San Gabriel Moun- eroding sedimentary deposits at the tains affinity, including Canterbury far southeast corner of the park, on LOCALLY RARE SPECIES bells (Phacelia minor), one of the soil probably deposited by the an- most abundant spring annuals in cient Los Angeles River that flows Several locally rare species were the park, becoming much more lo- nearby. Thousands of these plants found in substantial populations in calized farther west in the Santa bloom in early spring in the under- Griffith Park. Both Nevin’s brickle- Monica Mountains. story of open black walnut wood- bush (Brickellia nevinii) and Ft. The population of southern Cali- land. The tiny clay bindweed (Con- Tejon milk-aster (Stephanomeria fornia black walnut (Juglans califor- volvulus simulans) was found by a cichoriaceae) are frequent on rock nica) in the park is so large that it volunteer in 2010 on a moist, grassy outcrops and roadcuts throughout was not mapped during the survey. slope on heavy clay near the south- the park and even in surrounding Black walnuts form a continuous western border of the park just north residential areas to the south. East- woodland/high shrubland on sedi- of Hollywood, a site that also sup- wood’s manzanita (Arctostaphylos mentary soils in the southeastern ports the only park population of glandulosa ssp. mollis), chaparral pea corner of the park, and this is also a chocolate lily (Fritillaria biflora). (Pickeringia ), and interior dominant species in the chaparral Nevin’s barberry (Berberis nevinii) live oak (Quercus wislizenii var. fru- that cloaks the northern slope of the is arguably the most famous rare plant tescens) occur high atop Cahuenga park. in the park, and also the most prob- and Burbank peaks within a dense lematic. Most stands (dozens of chamise-manzanita chaparral above PATTERNS OF DIVERSITY plants each) occur in two popula- the 1,600-foot elevation level, with tions. Scattered plants located since smaller numbers of manzanita and a Although areas at the lowest el- 2007 have all been along major roads single clump of interior live oak near evations at the perimeter of the park (strongly suggesting a horticultural Mt. Hollywood. tended to support fewer target spe- origin), or just down-slope of estab- “Moss gardens,” where soil has cies, several featured a handful of lished populations. Whether plants collected at seeps on rock faces, sup- locally rare species. Important popu- at Griffith Park originated from lo- port a variety of bryophytes, ferns, lations, including the only known cally-collected seed decades ago— and rare annuals at Royce Canyon local occurrences, were found at possibly from now-extirpated native and east along the north side of Mt. multiple and scattered locations, populations in the Los Angeles area— Bell, including California saxifrage making it difficult to declare a given it seems likely that the current popu- (Saxifraga californica), Peninsular area unimportant for any rare plant. lation is not naturally-occurring onion (Allium peninsulare), Fremont Not surprisingly, the fewest target (USFWS 2009). Still, Griffith Park is star-lily (Zigadenus fremontii), and species overall were found in the within the historical range of the spe- (Royce Canyon only) a white-flow- northeastern corner of the park. This cies and all plants occur in a low- ered form of Cleveland’s shooting- area that has probably seen the most elevation, arid chaparral community star (Dodecatheon clevelandii). A few intensive human-caused impacts with a species composition very simi- areas of private land adjacent to the over the years, including the con- lar to known natural occurrences. park were found to be rich in scarce struction of a large landfill in the

22 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 1950s, the installation of a vast net- the distribution of rare and local- Nevin’s barberry, one of the rarest plants work of metal irrigation pipe de- ized plants can be an important first in California (in the wild), thrives in signed for fire control (but never step in a site’s conservation. The in- Griffith Park, although the entire popula- tion is suspected to have been introduced functional), and widespread plant- volvement of local volunteers not decades ago. It is a large shrub found very ing of eucalyptus and pines, ostensi- only contributes information to this locally on loose, gravelly soils within bly for “beautification” starting in process, it helps ensure that the hu- chaparral, and is endemic to southern the early 1900s. man community will be invested in California. Photographs by Gerry Hans. It also became clear in our survey resource conservation in “Holly- that most scrub-covered slopes in the wood’s backyard.” park were largely devoid of rare plants, with diversity mainly confined REFERENCES to microhabitats such as seeps in deep canyons, sandy/gravelly ridges, moss Brusha, R.F., ed. 2003. Native and ex- gardens, rock outcrops, and patches otic flora of Griffith Park: Source: Bill of grassland on wet clay soils. Areas Eckert. Unpublished checklist. of the park that combined these fea- California Native Plant Society. 2010. Inventory of Rare and Endangered tures predictably supported both a Plants. Vols. 7–10. http://cnps.site. high diversity of rare species and large aplus.net/cgi-bin/inv/inventory.cgi/ populations of them. Home. The identification and documen- Cartifact, Inc. 2007. Griffith Park tation of these microhabitats are (map). City of Los Angeles, Office critical to making management de- of Councilman Tom LaBonge. cisions that benefit biodiversity, and Consortium of California Herbaria

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 23 ABOVE: “Moss gardens,” or patches of moist soil within rock outcrops, are key areas for rare plants at Griffith Park. • LEFT: The prickly phlox (Leptodactylon californicum) occurs as scattered plants within the park, mainly in small openings in arid scrub on eroding ridges and slopes. Its bright pink flowers are unmistakable in spring. • BOTTOM LEFT: The striking Catalina mariposa lily (Calochortus catalinae) is one of three species of Calochortus in the park, and is found amid grasses on patches of heavy clay soil, especially those that stay moist through the spring.

(CCH). 2011. Data provided by the McAuley, M. 1985. Wildflowers of the participants of the Consortium of Santa Monica Mountains. Canyon California Herbaria. http://ucjeps. Publishing, Canoga Park, CA. berkeley.edu/consortium/. National Park Service (no date). Plant Cooper, D.S. 2010. Griffith Park Rare checklist for the Santa Monica Plant Survey. October 10, 2010. Un- Mountains National Recreation Area. published report. http://www.friends http://www.nps.gov/samo/nature ofgriffithpark.org. science/plants.htm. Cooper, D.S. Forthcoming. Flora of United States Fish and Wildlife Service. Griffith Park. Annotated checklist. 2009. 5-year review for Berberis Cooper, D.S. and P. Mathewson. 2009. nevinii (Nevin’s barberry). Carlsbad Griffith Park wildlife management Fish and Wildlife Office. August 14, plan. Report submitted to the Los 2009. Angeles Department of Recreation Wishner, C. 1997. Flora of the Santa and Parks by Cooper Ecological Monica Mountains: Synonymised Monitoring, Inc. http://www.griffith checklist and index. Crossosoma parkwildlife.org. 23(1):1-52. Gibson, A. and B.A. Prigge. 2003. Re- vised flora of the Santa Monica Daniel S. Cooper, Cooper Ecological Moni- Mountains. Final Report. Herbar- toring, Inc., 255 Satinwood Avenue, Oak ium, University of California, Los Park, CA 91377, dan@cooperecological. Angeles. com

24 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 A PLEA TO PROTECT WALKER RIDGE by Stephen W. Edwards

(Editor’s Note: The following article is an abbreviated version of a longer article that appeared in The Four Seasons, Journal of the Regional Parks Botanic Garden, Volume 14, Number 1, October 2011.)

alker Ridge, on the boundary of Lake and Colusa counties be- tween Clear Lake and Wthe town of Williams, forms an in- separable unit with Bear Valley, which adjoins the ridge to the east. The valley—thanks to the dedica- tion of generations of ranchers and timely action of conservation orga- nizations— displays in good years Map showing the proximity of Walker Ridge to Bear Valley. Source: Gregory Gallagher. by far the best remaining panora- mas of Northern California’s field park celebrated principally as a rem- That is why I have long felt that wildflowers. No other area comes nant of the wildflowers of the San Walker Ridge—most of which is close. Joaquin Valley as Thomas Jefferson under the jurisdiction of the Bureau Sediments on the valley floor are Mayfield and John Muir described of Land Management (BLM)—and strongly influenced by the magne- them (1850s and 1860s respec- parts of Bear Valley that might one sium-rich, calcium-poor ultramafic tively). So it is with Bear Valley and day be in public ownership, should rocks of Walker Ridge, and this al- Walker Ridge, which are to the origi- become a national monument. most certainly contributes to the nal, unspoiled Sacramento Valley of Walker Ridge preserves much of one wildflower displays. The ridge and the early 1800s what the Carrizo of California’s three greatest serpen- the valley have been a combined des- Plain National Monument is to the tine terrains (the others are The Ce- tination for wildflower lovers, re- San Joaquin Valley. dars in Sonoma County and the Trin- searchers, and university class study for decades. Visiting the one almost Green’s butterweed (Senecio greenei), an orange-hued serpentine endemic found on Walker always entails visiting the other be- Ridge, bears some of the most striking flower heads of any sunflower. It has stubbornly cause the contrast between their resisted all attempts by horticulturists to propagate it. Also pictured is common woolly sunflower (Eriophyllum lanatum). Photograph by Craig Thomsen. natural histories is fascinating. The valley with its vast, flowery panorama is made all the more enchanting by the bold backdrop of Walker Ridge.

OF NATIONAL PARK QUALITY

Walker Ridge and Bear Valley together are clearly of national park quality. The Carrizo Plain National Monument, which includes the plain as well as parts of the Temblor and Caliente ranges in Southern Califor- nia, is an excellent analogy. It is a

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 25 ABOVE: Serpentine scree slopes along the crest of Walker Ridge are precious preserves for rare, unusual, and beautiful plants. The proposal for a wind farm on the ridge includes an array of giant wind turbines, along with attendant roads, trenches, etc. Photograph by Phil Van Soelen, 2009. • RIGHT: Sky lupine (Lupinus nanus), creamcups (Platystemon californicus), purple owl’s clover (Castilleja exserta), and in the distance, tidy tips (Layia sp.), on the floor of Bear Valley, with Walker Ridge in the background, April 1995. Photograph by Stephen W. Edwards. ity ultramafic sheet around Mt. Eddy turbines, with all their associated in Siskiyou County), and these are equipment, facilities, pads, yards, three of the greatest serpentine ar- and roads, along the serpentine sum- eas on earth. Walker Ridge is a liv- mit of Walker Ridge. Those familiar ing classroom of inner coast range with the biology and scenery of natural history, from which one can Walker Ridge know that it is the last gaze eastward at Sierran snows and place in California that should be the Pleistocene volcanoes of the considered for any kind of intrusive Sutter Buttes, westward to the Clear development, and that is not merely Lake volcanic field, and north to the owing to its richness. Accessibility snow-capped peaks of Mendocino is another critical factor. Here one National Forest. can easily drive to many spectacular It makes sense to imagine Walker habitats, and the whole area is only Ridge and parts of Bear Valley as one to two hours by car from major California Serpentine NationaI population centers and universities. Monument. Serpentinite, which, owing to its fortunate chemistry, ac- fers many open serpentine scree counts for a greatly disproportion- UNPARALLELED RICHNESS, slopes decorated with striking flow- ate amount of high-quality Califor- BEAUTY ering gems that cannot endure com- nia wildflower habitat, has not re- petition on less severe sites. (Scree ceived adequate recognition at a fed- It has long been known that ser- is a gravely deposit that forms on eral (or state) level. Walker Ridge pentine meadows on Walker Ridge hillsides where bedrock is weather- offers an opportunity to rectify that are the place to go for great displays ing not far below the surface.) Many insufficiency and, at the same time, of flowers after Bear Valley has of these plants, too, can be enjoyed provide a park of surpassing beauty largely dried up. The ridge also of- in flower in summer. Road banks and fabulous biodiversity that could themselves, and natural benches be enjoyed by all. People make pilgrimages to Walker Ridge above them, are full of gorgeous to see the rare serpentine milkweed (Ascle- plants. Many of these plants may be pias solanoana). It can be found only on difficult for anyone with mobility sliding scree, through diligent searching. IMPENDING DEVELOPMENT Its low, ground-hugging habit makes it issues to find or reach away from THREAT intolerant of competition. Photograph by the road. Serpentine chaparral also Stephen W. Edwards. contains in its understory and rocky At the time of this writing, plans openings a host of exciting plants. are underway by a Canadian energy Streams that cut serpentine gorges company to develop an extensive along and near Bartlett Springs Road and elaborate array of giant wind cascade over travertine benches and

26 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 bring clear, cool water to lush and that botanists had driven by for de- Americans. Tuleyome, a conserva- extraordinarily diverse riparian veg- cades. tion organization centered in Wood- etation that includes other, equally Walker Ridge is one of the great- land, has been working hard to pre- beguiling wildflowers. The botani- est places left in California. Is this serve Walker Ridge as part of the cal richness of Walker Ridge is too where we would choose to build a proposed Berryessa-Snow Mountain great to enumerate. There are sev- conglomeration of huge (nearly 400 National Conservation Area. At the eral running plant lists, and they all feet tall) wind towers? Would we same time, the California Native keep growing as new discoveries are scrape off the serpentinite and rare Plant Society has formally proposed made. plant jewels for yards, pads, trenches, that the BLM make the ridge an Area Walker Ridge is not comprised and tarmac? Do we want to widen of Critical Environmental Concern. entirely of serpentinite. Much of the Walker Ridge Road for oversized I am proposing national monument southern half north of Highway 20 trucks, grade away its rich road status, but, whatever the name, the is on Great Valley Sequence sand- banks, replace tranquility and natu- critical thing is that the dignity of stones. These support a more com- ral beauty with fluster and rush, Walker Ridge be recognized with mon, less diverse chaparral and oversight, devastation, and weeds? the honor it deserves. woodland. Nevertheless, the huge Walker Ridge is rich in wild- fire of 2008 burned much of this life—foothill yellow-legged frogs Stephen W. Edwards, Regional Parks part of the ridge, creating the condi- seem to inhabit every serpentine Botanic Garden, c/o Tilden Regional Park, tions for sheets of wildflowers, and stream—and there is important ar- Berkeley, CA 94708-2396, sedwards@ exposing all kinds of lovely plants chaeological evidence of native ebparks.org

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 27 SERPENTINE SOILS AND WHY THEY LIMIT PLANT SURVIVAL AND GROWTH by Earl B. Alexander

erpentine evolves from pris- ROCKS FROM WHICH tine ultramafic rocks formed SERPENTINE SOILS beneath the oceans. Rocks DEVELOP which contain high levels of Siron and very high levels of magne- Serpentine is ecological jargon sium are alien on the continents, for ultramafic rock. The common where there is more calcium in the ultramafic rocks in California are rocks than magnesium. peridotite and serpentinite. Peridot- Serpentine soils are those that ite is the ultramafic rock produced develop from the weathering and by crystallization of magma beneath disintegration of ultramafic rocks. the oceans, and serpentinite is pro- They inherit much of their mineral duced by the hydrothermal alter- composition and chemistry from ation of peridotite. Serpentine was those rocks. Serpentine soils are so designated the state rock in Califor- different from others that relatively nia before geologists decided to call few plants are adapted for survival the rock serpentinite and restrict ser- in them. Ultramafic rocks and soils pentine to the name of the dominant cover only about 1% of the land area mineral in serpentinite (Wagner in California, yet 12% of the en- 1991). demic species in the state are re- The story of ultramafic rocks and stricted to serpentine soils (Safford serpentine soils begins in the mantle. et al. 2005). How are ultramafic The mantle is between the core and rocks and serpentine soils different the crust of Earth. Geologists be- from non-ultramafic ones, and why lieve that the composition of the are serpentine soils so difficult for upper mantle is comparable to that plants to cope with? of lherzolite, which is a type of peri- dotite. Below the oceans, partial melting in the upper mantle pro- Ophiolite, which consists of basalt, feeder dikes, gabbro, and harzburgite (peridotite). duces a magma that has more of the ELEMENT Serpentine forms by alteration of the elements with larger cations (posi- minerals (olivine and pyroxenes) in ABBREVIATIONS tive ions), such as potassium and peridotite. C carbon calcium, and less of the elements with smaller cations, such as mag- burgite, which is another type of Ca calcium nesium, than in lherzolite. peridotite. It is in harzburgite that Co cobalt Much of the magma rises the elements with smaller cations Cr chromium through the ocean crust at spread- such as magnesium, chromium, ing centers, or ridges, and flows out- manganese, iron, cobalt, and nickel H hydrogen ward under seawater, cooling rap- are concentrated following partial K potassium idly to form rounded blobs of lava melting of lherzolite. The sequence Mg magnesium that become a distinctive feature of of rocks from basalt, through a N nitrogen pillow basalt. Magma that does not swarm of dikes that have fed basalt rise to the top of the ocean crust upward, and gabbro to harzburgite, Ni nickel cools more slowly and forms gab- is called ophiolite. O oxygen bro, an intrusive igneous rock that Ophiolite is produced near the P phosphorous is the equivalent of basalt, an centers of large blocks of ocean crust extrusive igneous rock. Beneath the called plates. The ocean plates move S sulfur gabbro, residue from partial melting away from the spreading centers to- in the upper mantle forms harz- ward continental plates. Along the

28 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 way, the ophiolite cools, and sedi- sink to depths where they begin to terranes, and ophiolites have been ments accumulate on the ocean melt. Partial melting of the plates thrust over them. Granitic bodies floor. As the ophiolite cools, it be- and adjacent continental crust pro- called plutons, and volcanic depos- comes heavier. By the time spread- duces magmas that may rise to the its are commonly added to accreted ing ocean crust reaches a continent, surface and form volcanoes. Such terranes after the terranes are at- it is generally cool and heavy enough volcanoes that skirt the Pacific Ocean tached to a continent. The Sierra to sink under the edge of the conti- comprise a volcanic chain aptly Nevada batholith is an amalgamation nental plate into the mantle, a pro- called the ring of fire. The Cascade of many plutons, for example, and cess called subduction. Mountain Range from Lassen Peak volcanic rocks in the Coast Ranges, Subducted plates sink for many in Northern California to Mount such as those of Mount St. Helens, millions of years, and some ophiolite Baker in is a link in this postdate accretion of the Franciscan may reach depths of about 410 kilo- chain of volcanoes. complex. meters near the bottom of the upper The ultramafic rocks in sub- Most peridotite (mainly harz- mantle. Sediments are commonly ducted ophiolites must reach the burgite) that we see in California scraped off of the plates and accu- ground surface to produce serpen- has been at least partially altered to mulate in long trenches where the tine soils. Rocks can be raised to the serpentinite. The alteration, a pro- plates sink beneath continental mar- surface by faulting, or the rocks can cess called serpentinization, begins gins, with some sediments being be folded and rocks in the folds ex- under the ocean and continues even dragged under continental margins posed by erosion of the overlying after harzburgite has been incorpo- and plastered against the continen- rocks. The rated into the continental crust. tal crust, underplating it. Plates that are comprised largely of the Fran- Serpentinization is a low tempera- have lost their sedimentary cover ciscan complex of accreted oceanic ture process (< 550o C) in which

The Lassics barren on the boundary between Humboldt and Trinity Counties. Beyond the barren there is an open Jeffrey pine forest on moderately deep serpentine soils on the right, in contrast to a dense mixed conifer forest on nonserpentine soils on the left. All photographs by the author.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 29 LEFT: Travertine draped over the rocks and forming a small pool where alkaline water produced from the serpentinization of peridotite emerges from the ground, Sonoma County. • MIDDLE: Pillow basalt downstream from the outlet to Nicasio Reservoir, Marin County. • RIGHT: Serpentinization in an exhumed rock. water is added to the olivine and PLANT GROWTH ON concentrations are very high and Ca pyroxenes in harzburgite, and some SERPENTINE SOILS concentrations are very low in ser- calcium and silicon are lost from pentine soil parent materials, the Ca/ pyroxenes: Serpentine soils are those that Mg ratios are very low compared to have peridotite or serpentinite par- those in average continental crust harzburgite + water ===> ent materials. These two parent ma- and soils. serpentinite + calcium hydroxide + terials commonly produce soils with The very high Mg concentra- silica different physical properties, but the tions, relative to Ca, limit plant up- The water that reacts with harz- plants appear to respond more to take of Ca from soils. High Mg (and burgite generally lacks carbon diox- the chemical properties that are simi- low Ca) concentrations in plants in- ide; its main role is to provide oxy- lar for soils developed from either terfere with metabolic processes that gen and hydroxyl ions used in the parent material. Serpentine soils require Ca, excluding plants that formation of serpentine, which con- have a broad range of depths and have not developed mechanisms to tains more of these than the miner- water-holding capacities, as do soils deal with these Ca/Mg imbalances. als in harzburgite. Water left over with most other kinds of parent ma- If supplemental Ca is added to a from the serpentinization process terials (Alexander et al. 2007). serpentine soil, P becomes a plant contains calcium and is alkaline (pH Therefore, we can concentrate on growth-limiting element. Plants > 11). It is responsible for convert- the chemistry to learn why plants commonly respond favorably to the ing some of the gabbro and basalt in respond differently in ophiolites to calcium silicate (rodin- serpentine soils than A pictorial view of serpentinization. Water has entered along gite). Water from the serpentiniza- in other soils. fractures in peridotite and caused serpentinization, which tion of harzburgite that flows from The main ele- diminishes from the fractures toward cores that are only slightly serpentinized peridotite. the rocks to the ground surface re- ments required by acts with bicarbonates in meteoric plants are H, C, N, O, water (water that has gained carbon Mg, P, S, K, and Ca. dioxide in the atmosphere or soils) The first four of these to precipitate the calcium and form elements are in at- travertine. Note that carbon dioxide mospheric gases, or dissolved in water reacts with it to water, from which form carbonic acid that dissociates plants obtain them ei- into hydrogen (H+) and bicarbonate ther directly or indi- - (H(CO3) ) ions: rectly. Elements for which plants are de- + - Ca(OH)2 + H , H(CO3) ===> Ca pendent on the soil

(CO3) + 2H2O parent materials are Ca-hydroxide + carbonic acid ===> mainly Ca, Mg, K, travertine + water and P. Because Mg

30 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 accumulators. There are hundreds AID TO TERMINOLOGY of plant species that hyperaccumu- late Ni, and a few that hyperaccu- accreted – added to a continent mulate Co. There are only two Ni cations – positively charged ions hyperaccumulating species in Cali- gabbro – intrusive (produced below ground) equivalent of extrusive fornia, and both are in the mustard basalt () family: milkwort harzburgite – ultramafic rock produced from lherzolite (defined jewel flower ( polyga- below) beneath ocean crust loides) and two varieties of alpine pennycress (Noccaea montana) hyperaccumulators – plants that accumulate more than 0.1% by weight of nonessential elements (e.g., Ni or Co) (Reeves et al. 1983). The only Co hyperaccumulating species are lherzolite – ultramafic rock believed to reflect the composition of the found in Africa. upper mantle, lherzolite yields plutonic (intrusive) harzburgite, Plants that thrive on serpentine plus intrusive gabbro and extrusive basalt soils have unusual capabilities to uti- mafic lava – molten rock, or magma, that flows over the surface of the lize calcium when the alkaline earth earth and cools to form basalt, an aluminum silicate rock with elements are dominated by magne- large concentrations of alkaline earth cations (Ca and Mg) and iron sium, and to tolerate concentrations ophiolite – a section of the Earth’s oceanic crust and upper mantle of cobalt and nickel that are toxic that has been uplifted and exposed above sea level to most plants. Plants of intertidal peridotite – ultramafic rock formed by crystalli1zation of magma zones are also adapted to grow in under the ocean soils with low Ca/Mg ratios, but they pillow basalt – mafic lava that flows over the seafloor and cools to do not have to cope with potentially solidify in large blobs resembling pillows toxic concentrations of cobalt and pluton – a body of rock (see plutonic rock) formed at depth nickel. Although relatively few plants are adapted to grow in ser- plutonic rock – rock that crystallizes slowly from magma at great depths to form crystals visible without magnification pentine soils, many of them can- not compete with other plants on pyroxenes – magnesium silicate minerals, lacking aluminum nonserpentine soils and thus are serpentine – dominant mineral in serpentinite (and previously the endemics confined to serpentine name of the rock) soils. serpentinite – a greenish metamorphic rock consisting largely of serpentine REFERENCES subduction – sinking or pushing of an oceanic plate beneath another plate Alexander, E.B., R.G. Coleman, T. terrane – alien crustal block that has an origin different from that of Keeler-Wolf, and S. Harrison. 2007. adjacent rocks Serpentine Geoecology of Western North America. Oxford University trichomes – hairlike appendages to plant epidermal cells Press, New York. ultramafic rocks – rocks consisting mainly of magnesium silicate Reeves, R.D., R.M. Macfarlane, and minerals, with much iron substituting for some of the magnesium R.R. Brooks. 1983. Accumulation of vacuoles – spaces within cells that are filled with liquids nickel and zinc in western North American genera containing serpen- tine-tolorant species. American Jour- nal of Botany 70: 1297-1303. addition of N to serpentine soils, elevated concentrations of these el- Safford, H.D, J.H. Viers, and S.P. but the same is also true for plants ements. They may selectively ex- Harrison. 2005. Serpentine ende- growing in other kinds of soils. clude the elements from root up- mism in the California flora: A data- The elements that are much take, confine them to roots, or accu- base of serpentine affinity. Madroño more concentrated in serpentine mulate them as chelated forms or 52: 222-257. Wagner, D.L. 1991. The state rock of soils than in others may be toxic to precipitates in the vacuoles of epi- California: Serpentine or serpenti- plants. These elements are Cr, Ni, dermal cells in leaves or concentrate nite? California Geology 44: 164. and Co. Some plants such as Sargent them in trichomes on the leaves. cypress (Cupressus sargentii) and Plants that accumulate hundreds Earl Alexander, Soils and Geoecology, leatheroak (Quercus durata) have de- of times more of an element than 1714 Kasba Street, Concord, CA 94518, veloped mechanisms to deal with most other plants are called hyper- [email protected]

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 31 SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA by Hugh D. Safford

ltramafic “serpentine” soils stricted in their range to California; area of approximately 2,300 square and rock outcrops support ultramafic endemics are only found miles in California (Harrison et al. an exceptionally high num- on ultramafic soils. About 11% of 2000), which makes up around 1.5% ber of endemic plant taxa the taxa found on the California Rare of the state’s area. However, as Earl Uin California, far out of proportion Plant Rank 1B (formerly CNPS List Alexander notes in the accompany- to their limited representation on 1B, critically rare taxa) are either ing article, inexactitude in mapping the California landscape. California strictly or broadly endemic to ultra- may inflate that area somewhat. “endemic” plant taxa are species, mafic substrates. Rare native taxa subspecies, and varieties that are re- that are not serpentine endemics also find important habitat footholds on NUMBERS AND TYPES OF ultramafic substrates, where they can SERPENTINE PLANTS escape competition from more ag- gressive, often non-native species. Serpentine endemism is a key component of the diversity of the California flora (Raven and Axelrod WHAT ARE SERPENTINE 1978, Kruckeberg 2002, Safford et SUBSTRATES? al. 2005). In 2005, we carried out an assessment of serpentine endemism By definition, ultramafic rocks in the California flora (Safford et al. are igneous and metamorphic rocks 2005). We found that of the 1,416 that contain less than 45% by weight endemic species listed in The Jepson

silica (SiO2) or, alternatively, sup- Manual, 1st edition (Hickman 1993), port a mineral content that is greater about 176 could be ranked as either than 90% “mafic” (i.e., magnesium “strict” or “broad” serpentine en- and iron silicate minerals like oliv- demics (“strict” having over 95% of ine, pyroxenes, or hornblendes) its occurrences on serpentine soils (Ehlers and Blatt 1982). Ultramafic and “broad” having over 85%). This rocks originate primarily in the constituted about 12.7% of the Cali- earth’s mantle, and find their way to fornia endemic flora. the surface by way of plate tectonics Since the 2005 analysis, many and mountain building, and subse- changes have occurred in the tax- quent erosion of overlying geologic onomy and classification of the Cali- layers. fornia flora, and a surprising num- Ultramafic rocks and the soils ber of new species have been de- that develop on them (often collo- scribed. Information from the Con- quially called “serpentine” after their sortium of California Herbaria most important mineral constitu- (CCH, http://ucjeps.berkeley.edu/con- ents) are characterized by critically sortium/) records, The Jepson Manual, low levels of most principal plant 2nd edition (Baldwin et al. 2012), nutrients (nitrogen, phosphorus, and the scientific literature was used potassium, calcium), exceptionally to decide how to rank newly de- high levels of magnesium and iron, scribed, combined, or split taxa with and a suite of highly toxic trace ele- respect to their preference for ser- ments including chromium, nickel, pentine soils. and cobalt (Alexander et al. 2007). This new information shows that In all, ultramafic rocks occupy an approximately 243 taxa are endemic

Some of the more striking serpentine forbs include (TOP TO BOTTOM): Indian Valley brodiaea (Brodiaea coronaria ssp. rosea), San Luis mariposa lily (Calochortus obispoensis), and talus fritillary (Fritillaria falcata). All are restricted to a handful of outcrops in the North or South Coast ranges. Photographs by Rick York, Bob Case, and John Game.

32 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 to ultramafic substrates in Califor- TABLE 1. NUMBERS OF SERPENTINE ENDEMIC AND nia, including 176 full species. These NEAR ENDEMIC TAXA FOR PLANT FAMILIES WITH AT numbers are very close to our 2005 LEAST THREE ENDEMIC TAXA ON ULTRAMAFIC numbers (246 taxa, 176 full spe- SUBSTRATES IN CALIFORNIA cies), although some of the names have changed, some taxa were FAMILY SERPENTINE AFFINITY SCORE dropped from the list, and some new taxa were added. In addition, due to Strict Broad Near Total changes in and two de- endemics endemics endemics taxa1 cades of additional field collection, the estimated number of plant spe- 26 11 8 105 cies endemic to California has actu- Brassicaceae 19 3 4 37 ally dropped to 1,315 (Baldwin et Polygonaceae 11 8 3 41 al. 2012), so the contribution of ser- pentine endemism to the California 6 6 3 37 endemic flora has risen to 13.4%! Apiaceae 7 3 3 32 Table 1 provides a summary of ser- Orobanchaceae 7 3 0 19 pentine endemic richness in Cali- fornia. Altogether, the families in Linaceae 8 1 0 14 Table 1 contribute almost 90% (216 Ericaceae 5 3 2 15 of 243) of the total serpentine en- Alliaceae 4 3 4 24 demic diversity in California. Asteraceae is the top family in 4 3 3 24 all categories listed, while second Polemoniaceae 6 1 1 18 and third place are held by Brassi- Boraginaceae 6 0 5 24 caceae and Polygonaceae. Figure 1 plots the families listed in Table 1. Caryophyllaceae 5 1 3 18 The first eight families include more Lamiaceae 4 2 3 17 than 50% of the taxa in my endemic Campanulaceae 3 2 3 13 database. Asteraceae includes about 15% of the serpentine endemic taxa Crassulaceae 5 0 2 13 in the list, a similar proportion to its Cyperaceae 2 3 2 11 contribution to the total California Onagraceae 3 2 2 13 endemic flora. Five of the next six families (all but Orobanchaceae) are Rhamnaceae 4 1 1 15 substantially overrepresented in the Agavaceae 3 1 0 6 serpentine flora, which means they Convolvulaceae 1 3 1 6 have been important evolutionary sources of new species on serpen- Rubiaceae 3 1 0 8 tine soils. Themidaceae 2 2 3 10 The most diverse genera of ser- Montiaceae 0 3 2 16 pentine endemism are listed in Table 2, topped by Eriogonum and Streptan- Plantaginaceae 1 2 2 15 thus, and followed by Hesperolinon Poaceae 1 2 0 18 and Arctostaphylos. The genera in Totals (all families) 164 79 72 661 Table 2 include more than 50% of the total taxa endemic to serpen- Familes are ordered by their relative numbers of serpentine endemics (strict plus tine soils in California. Basically, broad endemics). Strict endemics have >95% of their documented occurrences most of the serpentine endemic plant on ultramafic substrates; broad endemics (>85%-94.9%); near endemics (>75%- diversity in California is driven by 84.9%). The latter group is the transitional group from “strong indicators” to only a few dozen genera (in only a broad endemics in Safford et al. (2005). few dozen families) that have evolved many species on serpentine (and 1 Total of all taxa in our database, from strict endemics to indifferent taxa. sometimes other harsh) substrates. Probably the flagship for serpen-

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 33 tine endemic plants in FIGURE 1. CALIFORNIA PLANT FAMILIES WITH AT LEAST THREE SERPENTINE ENDEMIC California is the genus TAXA, AS A PERCENTAGE OF THE TOTAL CALIFORNIA SERPENTINE ENDEMIC FLORA Streptanthus (Brassica- ceae), which includes at least 16 plant species or subspecies that are strictly or broadly endemic to ser- pentine soils. Streptanthus (commonly called the “jewelflowers”) is an ex- ample of a neoendemic taxon, a relatively recently evolved plant group that continues to actively gen- erate new species. Ex- amples include Socrates mine jewelflower (Strep- tanthus brachiatus), which is restricted to a few ser- pentine locations at the junction of Lake, Napa, and Sonoma counties, and milkwort jewelflower (Strep- TABLE 2. GENERA WITH AT LEAST FOUR SERPENTINE tanthus polygaloides), a species from ENDEMIC TAXA IN CALIFORNIA the Sierra Nevada which has evolved the rare ability to hyperac-cumulate Genus Family Endemic taxa nickel, a highly toxic heavy metal Eriogonum Polygonaceae 16 that is usually deadly in the concen- Streptanthus Brassicaceae 16 trations found in the plant. Of the 243 serpentine endemics Hesperolinon Linaceae 9 identified in the updated assessment, Arctostaphylos Ericaceae 8 86% are forbs (210 out of 243), with one-third of those being annuals and Allium Alliaceae 7 two-thirds perennials (Figure 2). Lomatium Apiaceae 7 Some of the more charismatic forbs are in the old lily family, now split Packera Asteraceae 6 into numerous families. Examples Calochortus Liliaceae 5 pictured are Indian Valley brodiaea Carex Cyperaceae 5 (Brodiaea rosea), San Luis mariposa lily (Calochortus obispoensis), and Cordylanthus Orobanchaceae 5 talus fritillary (Fritillaria falcata), all Calystegia Convolvulaceae 4 of which are restricted to a handful of ultramafic outcrops in the North Castilleja Orobanchaceae 4 or South Coast Ranges. Cirsium Asteraceae 4 Graminoids (grasses, sedges, and Erigeron Asteraceae 4 rushes), shrubs, and trees comprise only 14% of the endemic serpentine Fritillaria Liliaceae 4 flora (Fig. 2). Leather oak (Quercus Galium Rubiaceae 4 durata var. durata) and musk brush (Ceanothus jepsonii) are serpentine Harmonia Asteraceae 4 endemic shrubs that often grow to- Lessingia Asteraceae 4 gether in the North Coast Ranges. One of the few tree taxa that is a Minuartia Caryophyllaceae 4 serpentine endemic is Sargent cy- Monardella Lamiaceae 4 press (Hesperocyparis sargentii), Phacelia Boraginaceae 4 which is found from Mendocino

34 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 County to Santa Barbara County, mostly at sites with moister soils or periodic summer fog. Since most ultramafic outcrops are very limited in size, many ser- pentine endemic plant taxa also have small geographic ranges. Combined with the occurrence of many ser- pentine outcrops in populated (and growing) coastal and foothill urban and suburban areas, this has led to an overrepresentation of serpentine endemic plants among the state’s endangered, rare, and sensitive plant taxa (CNPS Inventory of Rare and Endangered Plants, http://www.rare plants.cnps.org/). The conservation status of the taxa in my database has not yet been updated, but in 2005, 186 (77%) of ceanothus (Ceanothus ferrisiae), the taxa in the endemic category found only in the low mountains (strict plus broad endemics) were southeast of San Jose; and Kellogg’s included in the CNPS Inventory of buckwheat (Eriogonum kelloggii), Rare and Endangered Plants (Safford which grows only at the top of Red et al. 2005). California Rare Plant Mountain in Mendocino County. Rank 1B, which includes federal and state threatened and endan- gered taxa, as well as taxa that CNPS WHERE SERPENTINE believes warrant listing due to their ENDEMICS ARE FOUND rarity, included 105 serpentine en- demics. Harrison et al. (2006) carried out Serpentine endemics listed as a detailed statistical analysis of the federal or California endangered factors influencing the geographic include Tiburon Indian paintbrush distribution of serpentine endemism (Castilleja affinis ssp. neglecta), in California. They found that local which is restricted to a couple of richness (the number of serpentine All three of these serpen- ultramafic outcrops in the north endemic plants) is primarily driven tine endemic species are ; coyote by an interaction between the re- listed as federal or Cali- fornia endangered. TOP TO BOTTOM: Coyote ceano- FIGURE 2. LIFEFORMS OF SERPENTINE ENDEMIC PLANT TAXA IN CALIFORNIA thus (Ceanothus ferrisiae), Kellogg’s buckwheat (Erio- gonum kelloggii), Tiburon Indian paintbrush (Cas- tilleja affinis ssp. neglec- ta). Photographs by Neal Kramer, Steve Edwards, and Kaylea Eickhoff.

gional richness of en- demics (i.e., there are more species to choose from in a diverse re- gion) and local environmental fac- tors such as overstory (shrub and tree) cover, litter cover, the rocki- ness of the soil, and soil chemistry.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 35 The greatest number of serpentine endemics in California are to be found in the Klamath and North Coast Ranges. ABOVE: Flame ragwort (Packera greenei), one of the few orange- flowered members of the sunflower family. Photograph byDick O’Donnell. • LEFT: Siskiyou inside-out flower (Vancouveria chrysantha), a shrub. Photograph by David McClurg.

precisely because the harsh rocky nifers are those whose cones only substrates allowed them to escape open and release seed at high tem- competition (Brooks 1987). At the peratures, usually during fires.) same time, some ultramafic sites Figure 3 portrays the distribu- are too harsh even for serpentine tion of serpentine endemic taxa by endemics (Harrison et al. 2006). Due geographic region. The center of ser- to their low soil nutrient status, ul- pentine endemism in the California tramafic outcrops tend to support Floristic Province is in the North relatively slow growing vegetation Coast and Klamath Ranges of north- and low levels of plant biomass. In western California (and southwest- California, with its Mediterranean- ern , not included here). In like climate and frequent fires, ul- addition, most serpentine endemics tramafic outcrops tend to burn less are found in habitats below 1500 Localities that have rockier soils have often than surrounding, more fer- meters (5,000 feet) (Figure 4). higher richness of endemics, while tile substrates (Safford and Harrison At the regional scale, Harrison overstory cover, litter cover, and the 2004). Thus serpentine areas may et al. (2006) found that serpentine relative balance of soil magnesium provide refuge from both biotic endemic richness is best explained versus calcium have curvilinear competition and ecological distur- by a model including regional pre- (“hump-shaped”) relationships to bances like fire (Safford and Mallek cipitation and regional vegetation local endemic richness, such that 2010). The abundance of serotinous productivity, the area of ultramafic high and low values of these vari- conifers like knobcone pine (Pinus rock in the region, and how long the ables support less richness and in- attenuata, not an endemic) and ultramafic rocks have been avail- termediate values support higher Macnab cypress (Hesperocyparis mac- able for plant colonization. Regions richness. nabiana, a broad endemic) on ser- with higher endemic diversity were Many, if not most, serpentine pentine soils is probably primarily those with higher precipitation and endemics are confined to ultrama- due to the lower frequencies of fire (regional) productivity, more ultra- fic habitats (Harrison et al. 2006) in these locations. (Serotinous co- mafic rock, and longer availability

36 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 of ultramafic substrates for coloni- FIGURE 3. GEOGRAPHIC DISTRIBUTION OF SERPENTINE ENDEMIC TAXA IN CALIFORNIA zation. This helps to explain the pat- terns in Figs. 3 and 4, as the Kla- math and North Coast Ranges are the wettest part of California, and support the most extensive ultra- mafic outcrops and oldest serpen- tine soils in the state. Examples of interesting Klamath and North Coast Range serpentine endemics include serpentine milkweed (Asclepias sola- noana), surely one of the strangest looking plants in the California flora; flame ragwort (Packera greenei), oc- casionally found on nonserpentine soils and one of the few orange- flowered members of the sunflower family; and Siskiyou inside-out “Total endemics” includes all Califiornia serpentine endemic taxa present in a given region. “Restricted flower (Vancouveria chrysantha), a endemics” includes only those taxa restricted to a given region. shrub whose strange flowers re- semble the shooting star. FIGURE 4. ELEVATIONAL DISTRIBUTION OF SERPENTINE ENDEMIC TAXA IN CALIFORNIA, Reduced richness of endemics at AS WELL AS ALL TAXA IN THE SERPENTINE AFFINITY DATABASE higher elevations is due principally to the relatively limited areas of ul- tramafic rocks above 1,500 meters, but probably also to reduced pro- ductivity connected to lower tem- peratures and a shorter growing sea- son. That said, there are some high elevation serpentine endemics in California, including Trinity buck- wheat (Eriogonum alpinum), and Mt. Eddy draba (Draba carnosula), both found about 2,100 meters (7,000 feet) in the Scott Mountains west of Mt. Shasta. In summary, the “average” Cali- fornia serpentine endemic species is a dicotyledonous, perennial forb from a short list of plant families and genera, rare enough to be listed Taxa are counted in every elevational belt where they occur. as warranting conservation concern, and restricted (primarily) to ultra- available for long enough periods of that is projected for the 21st century mafic soils because the difficult en- time to provide for the evolution of is especially likely to impact plants vironment filters out faster-growing a large species pool of serpentine- whose habitat is restricted to dis- competitors. The highest levels of adapted plant taxa. junct patches of soil scattered across endemic richness on ultramafic sub- Recent research on the effects of the landscape. It will be expensive strates in California are found on current and projected future climatic and both logistically and politically relatively harsh (although not ex- trends on narrowly distributed en- difficult to conserve serpentine en- treme) soils in areas of relatively demic plants (Damschen et al. 2010) demic taxa on a species-by-species high precipitation and vegetation suggests that we should be worried basis. Conservation of the remain- productivity, at low to moderate el- about the long-term prospects for ing significant areas of unprotected evations, in regions where large ar- many of California’s serpentine serpentine habitat, especially in and eas of ultramafic rocks have been endemics. The extreme warming around urban and suburban areas in

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 37 The genus Streptanthus and genus Eriogonum top the list of those containing the greatest number of serpentine endemic species in California. Pictured are (CLOCKWISE FROM TOP LEFT): Socrates mine jewelflower (Streptanthus brachiatus), milk- wort jewelflower (Streptanthus polygaloides), and Trinity buck- wheat (Eriogonum alpinum). Photographs by Dick O’Donnell, Bob Case, and Julie Kierstead Nelson. the Bay Area, southern North Coast Ranges, and Sierra Nevada foothills, would be a good—although certainly not sufficient— alternative.

REFERENCES Alexander, E.B., R.G. Coleman, T. Keeler-Wolf, and S. ing Robert H. Whittaker’s Siskiyou J.H. Viers, and K.F. Davies. 2006. Re- Harrison. 2007. Serpentine Geoecol- sites (Oregon, USA). Ecology 91: gional and local species richness in ogy of Western North America. Ox- 3609-3619. an insular environment: Serpentine ford University Press, New York, NY. Ehlers, E.G., and H. Blatt. 1982. Petrol- plants in California. Ecological Mono- Baldwin, B.G., D.H. Goldman, D.J. Keil, ogy. W.H. Freeman and Company, graphs 76: 41-56. R. Patterson, T.J. Rosatti, and D.H. San Francisco, CA. Hickman, J.C., ed. 1993. The Jepson Wilken, eds. 2012. The Jepson Grime, J.P. 2001. Plant Strategies, Veg- Manual: Higher Plants of California. Manual: Vascular plants of California. etation Processes, and Ecosystem University of California Press, Ber- 2d edition. University of California Properties. 2d ed. Wiley, Chichester, keley, CA. Press, Berkeley, CA. UK. Jaffré, T. 1992. Floristic and ecological Brooks, R.R. 1987. Serpentine and its Harrison, S.P., J.H. Viers, and J.F. diversity of the vegetation on ultra- Vegetation. Dioscorides Press, Port- Quinn. 2000. Climatic and spatial mafic rocks in New Caledonia. In land, OR. patterns of diversity in the serpen- The Vegetation of Ultramafic (Serpen- Damschen, E.I., S. Harrison, and J.B. tine plants of California. Diversity tine) Soils: Proceedings of the First In- Grace. 2010. Climate change effects and Distributions 6: 153-161. ternational Conference on Serpentine on an endemic-rich flora: Resurvey- Harrison, S.P., H.D. Safford, J. Grace, Ecology, eds. Baker, A.J.M., J. Proc-

38 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 TOP: Mt. Eddy draba (Draba carnosula) is among the comparatively few serpentine endemics found above 7,000 feet. Photo- graph by Steve Matson. • BOTTOM LEFT: Stebbins’ tarweed (Harmonia stebbinsii) is an endangered species found on shallow, rocky soils, and is on California Rare Plant Rank 1B.2 (formerly CNPS List 1B.2). Photograph by Julie Kierstead Nelson. BOTTOM RIGHT: There are three varieties of fountain thistle found in and around seeps and streams. All are strict serpentine en- demics. This is Cirsium fontinale var. cam- pylon. Photograph by Bob Case. tor, and R.D. Reeves, 101-107. In- tercept Ltd., Andover, UK. The Evolution and Ecology of a Model Kruckeberg, A.R. 1984. California System, eds. S.P. Harrison and N. Serpentines: Flora, Vegetation, Ge- Rajakaruna, 249-274. University of ology, Soils and Management Prob- California Press, Berkeley, CA. lems. University of California Press, Reeves, R.D., A.J.M. Baker, A. Borhidi, Safford, H.D., J.H. Viers, and S. Berkeley, CA. and R. Berazain. 1999. Nickel hyper- Harrison. 2005. Serpentine ende- Kruckeberg, A.R. 2002. Geology and accumulation in the serpentine flora mism in the California flora: A data- Plant Life: The Effects of Landforms of Cuba. Annals of Botany 83: 29-38. base of serpentine affinity. Madroño and Rock Types on Plants. University Safford, H.D., and S. Harrison. 2004. 52: 222-257. of Washington Press, Seattle, WA. Fire effects on plant diversity in ser- Raven, P.H., and D.I. Axelrod. 1978. Ori- pentine vs. nonserpentine chaparral. gins and relationships of the Califor- Ecology 85: 539-48. Hugh Safford, USDA Forest Service, Pa- nia flora. University of California Pub- Safford, H.D., and C.R. Mallek. 2010. cific Southwest Region, 1323 Club Drive, lications in Botany, Vol. 72. University Disturbance and diversity in low pro- Vallejo, CA 94592, hughsafford@fs. of California Press, Berkeley, CA. ductivity ecosystems. In Serpentine: fed.us

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 39 The Joshua tree (Yucca brevifolia) is the ubiquitous tree of the Mojave Desert. This specimen is growing in Red Rock Canyon State Park in the eastern Mojave next to fourwing saltbush (Atriplex canescens). The Mojave is home to many plant galls, including some of the newly discovered ones mentioned in this article. All photographs by the author except as noted. PLANT GALLS: DESERT TREASURES by Ron Russo

n 1962 a colleague and I drove to There they were within the pe- the New World to me, full of wild the Mojave Desert from Sacra- ripheral glow of the headlights, alien creations and countless new discov- mento for our first botanical col- forms that turned out to be Joshua eries. lecting trip to this area. Sometime trees (Yucca brevifolia), Spanish Even though nearly 50 years Iin the middle of the night, as we bayonets (Yucca baccata), and blad- have passed and I now feel comfort- were driving downslope into north- der pods (Isomeris arborea) dotting ably familiar with many of the plants ern reaches of the desert, I woke up the upper elevations of the Mojave of the Mojave and Sonoran Deserts, from a deep sleep in the passenger Desert. As it was to the earliest ex- I still feel the excitement of ventur- seat to see the strangest looking plorers when they landed on the ing into the “New World” each time plants I had ever seen. shores of North America, this was I enter this unique ecosystem. And

40 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 with each trip, I encounter new spe- brush, (C. nause- cies that only stir the imagination of osus) in the north- what is yet to be discovered. Over ern Mojave. Later, I these many years, I have made hun- found the same gall dreds of pilgrimages to these deserts insect on yellow rab- in search of plant galls—tumor-like bitbrush (C. viscidi- growths on plants induced by in- florus), closer to our vading organisms to house their lar- home at the time in vae, in the case of insects—and to Mount Shasta, which study seasonal changes to the plants allowed me to make that host them. weekly visits over a Plant galls are specific in size, period of several shape, and color to each species of months and collect insect that induces them. Further- the galls at just the more, gall insects have evolved right moment to strictly dependent on either a genus successfully rear the of plants or a species within. A new adult midges. These species of gall midge, for example, were later identified was discovered in 2004 on desert as a new species of holly (Atriplex hymenelytra) that does midge in the genus, not occur on any other host plant. Rhopalomyia. Rab- The exact mechanism(s) responsible bitbrush also plays for species-specific designs in plant host to one of the galls and species-specific host selec- rarest and strangest interrelation- tion has yet to be uncovered. To ships in the insect world where the date, I have found 101 species of gall midge (Rhopalomyia bigelo- these gall-inducing insects that are viae) induces a gall in the gall of the dependent on desert plants during bubble gall tephritid (Aciurina trixa). their lifecycles. And of these, 38 were These “endogalls” as they are called, previously unknown to science. represent the ultimate to me in Some shrubs support relatively micro-niche evolution. large numbers of gall insects. For Another strange occurrence in- example, at least 16 species of gall volves Cooper’s boxthorn (Lycium midges are found only on the leaves cooperi). I had been studying a par- and flowers of creosote bush (Larrea ticular small group of plants near tridentata). The creosote stem gall Halloran Pass off Highway 15 east midge (Asphondylia auripila) in- of the small desert community of duces the largest gall, about the size Baker. For three years I had col- of a golf ball, on these resilient lected elliptical stem galls belonging shrubs, and it was ground and to a new species of moth, trying to TOP: Desert holly (Atriplex hymenelytra) is smoked by the Sere Indians of the rear the adults (later identified as widespread across the deserts into Southwest. Finding other species, Gnorimoschema sp.). During these and New Mexico. Its normal white color is offset in spring when large, bright pink, however, like the galls of the creo- three years, the stem galls were the cotton-candy-like galls develop from some sote antler gall midge (A. digitata) only galls found on the specific of its buds. • BOTTOM: Creosote bush (Larrea that look like tiny moose antlers, or shrubs being examined quarterly. tridentata) is favored by many plant galls. the galls of the leaf club gall midge Then, seemingly out of nowhere, (A. pilosa), takes careful and disci- dozens of cabbage-like bud galls ap- Several rather innocuous look- plined observation. peared one spring. While I could ing shrubs also support gall insects. My next favorite among desert not locate shrubs nearby from which The relatively unassuming burro- hosts is rabbitbrush (Chrysothamnus they may have spread to these new weed and related ragweeds (Ambro- spp.), which is host to 15 species of hosts, they certainly must have been sia spp.) support at least seven spe- gall insects, including a few new in the region. It took another year cies of gall midges, while the vari- species. One such new species, found for me to rear the adults and have ous saltbushes and shadscales (Atri- in 2004, induces a cotton-like ter- them identified as a new midge spe- plex spp.) support eight species. minal flower gall on rubber rabbit- cies in the genus Contarinia. Other desert plants that also sup-

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 41 port gall insects include: bladder sage (Salazaria mexicana), indigo bush (Psorothamnus arborescens), golden- bush (Ericameria cuneata), cheese- bush (Hymenoclea salsola), desert broom (Baccharis sarothroides), brittlebush (Encelia spp.), Mormon tea (Ephedra spp.), horsebrush (Tetradymia spp.), and catclaw (Aca- cia greggii). In 2009, a midge was reared from a bristly tubular gall on catclaw that, according to Dr. Raymond Gagné, a leading gall midge taxonomist, was so astonishingly different from all other known species, it would likely require the description of a new ge- nus and, perhaps, a new family. How many other great surprises are out there? But of all the interesting desert plants that have drawn my atten- tion, it is the desert oaks that have dominated much of my field time these last several years. Muller’s oaks (Quercus cornelius-mulleri) and Pal- mer’s oaks (Q. palmeri) have become a fount of new discoveries. Not only do they share species more common to their northern relatives, but they also host several recently discovered ABOVE: At least 16 species of gall midges species. (flies) are found only on the creosote bush, including Asphondylia auripila, which While there is a small cluster of induces a gall about the size of a golf ball. Muller’s oaks in Joshua Tree Na- This midge occurs throughout the range tional Monument, a much larger and of creosote bush in the southwest. This contiguous stand of these oaks oc- gall was ground and smoked by the Sere curs in the Northern Santa Rosa Indians of . LEFT: The gall Asphon- dylia digitata, often described as looking Mountains just southwest of Palm like moose antlers, is commonly found on Desert. These trees support several creosote bush. • BELOW LEFT AND RIGHT: The species of cynipid wasps including tiny creosote stem gall midge (about .15 the beaked twig gall wasp (Dishol- inch), and an equally small wasp (about caspis plumbella), and the saucer gall .18 inch) emerging from a beaked twig gall. Both insect photographs by Peter J. wasp (Andricus gigas), more com- Bryant. monly associated with related white oaks of the Central Valley and sur- rounding foothills. Muller’s oaks, however, have yielded several new species not seen elsewhere. Some of these may prove to be the elusive and currently un- known alternate generations of known species. Specifically, cynipid wasps exhibit a rare phenomenon where a spring bisexual generation alternates with a summer-fall gen-

42 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 TOP LEFT: Rhopalomyia sp. on rabbitbrush. This unique flower gall, a new species of Rhopalomyia, was discovered on two different species of rabbitbrush, one in the northern Mojave and the other in Northern California. • ABOVE LEFT: One of the strangest interrelationships in the Mojave has one insect inducing a gall in the gall produced by another insect. Pictured here is Aciurina trixa on rabbitbrush with the endogall of Rhopalomyia bigeloviae protruding. • ABOVE RIGHT: Cooper’s boxthorn, Lycium cooperi, is widespread throughout the desert’s sandy and rocky flats and washes east into Utah and Arizona. This shrub also supports a new species of stem-galling moth, Gnorimoschema sp. eration of females only. Many spe- to science. Such is the mystery that far, all of which I have reared for cies are currently known only by surrounds what I call the melon gall, identification. the adults from one generation, the lobe gall, and the lemon gall Even though Palmer’s oaks are while others may be completely new found only on Muller’s oaks thus closely related to canyon live oak

New gall species encountered by the author. LEFT: Contarinia sp. on Cooper’s boxthorn (Lycium cooperi). The gall, which looks like a head of cabbage, was found in the eastern Mojave. • MIDDLE: Rhopalomyia sp. on desert broom (Baccharis sarothroides). The galls of this new species of Rhopalomyia are heavily parasitized (94%) by torymid wasps in one area in the northeastern Mojave. • RIGHT: New gall species on catclaw (Acacia greggii). The life cycle of this midge and many other gall insects in the desert is controlled by major rain events that influence the development of new buds.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 43 (Quercus chrysolepis) and huckle- perhaps unanswerable question of berry oak (Q. vaccinifolia) and share how many other species might be some of their gall associates, Palmer’s discovered with a concentrated, oaks support an interesting group long-term effort. The notion that of largely unknown new species, Palmer’s oaks, Muller’s oaks, and unique to them. While five species desert scrub oak (Q. turbinella) as have been reported on Palmer’s oaks, well as other desert shrubs and trees five additional new species were dis- could support dozens of new spe- covered just in 2009. cies, as well as unknown alternate What drives my seemingly end- generations, is the magnet that con- less quest for new plant galls is the tinues to lure me back. My work in the desert over all these years documenting the occur- rence of gall-inducing insects and mites has not been what I would call earth-shattering, but it does reveal a tiny facet of desert life not previ- ously well understood. When you consider a shrub like rabbitbrush or creosote bush and take into account all of the associate species depen- dent on it—the pollinators, leaf-eat- ing caterpillars, leaf miners, nectar thieves, and the complex mix of sev- eral species of gall insects—you have a biological universe within a single shrub. Then when one considers the ramifications of the presence of a single species of gall insect with all of its predators, parasites, their para- sites, and the insects that eat gall tissue exclusively, the importance of these species in the grand web of desert life staggers the imagination. By recognizing this tiniest of threads, we might just better understand the whole fabric of desert life. Clearly, the biological diversity and the na- ture of interrelationships in the desert are far more complex than we currently understand. And it is this complexity that makes the risk of losing species and systems that we don’t even know exist that much more critical and irreversible. TOP: The beaked twig wasp gall (Disholcaspis plumbella) occurs on scrub oak (Quercus A couple of years ago I found berberidifolia) elsewhere in the state, but also occurs on Muller’s oak (Q. cornelius-mulleri) myself standing next to a Palmer’s in the desert. • MIDDLE LEFT: New species of bud gall on Muller’s oak. The galls of this new species produce males and females in the late spring, which suggests there is an alternate oak, at the head of a canyon in late generation later in the summer consisting only of females. • MIDDLE RIGHT: The lobe gall, March, buffeted by a stiff, cold wind, a new species, was first found in the northern Santa Rosa Mountains above Palm Desert chilled and hungry, wondering what on Muller’s oak in 2009. Normal anthocyanin pigments found in plant tissue may be I was doing there. And then a small, concentrated in specific areas, as in certain galls, where the red-orange is expressed. How and why this happens and what role the gall insect plays remain a mystery. Photograph green, lumpy gall I had never seen by Joyce Gross. • BOTTOM: Plant gall seekers are continually rewarded with new finds, such before caught my eye. After all these as this small, green, “lumpy” gall discovered by the author in 2010. years, the Mojave and Sonoran

44 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 THE PALMER’S OAK MYSTERY ne of the grand mysteries that has puzzled me for years, until Ojust recently, is the occurrence of the same cynipid gall wasps on Palmer’s oaks that are widely separated by hundreds of miles. I examined three Palmer’s oaks, for example, at the head of Short Canyon in the Owen’s Peak Wilderness on the eastern side of the southern Sierra Nevada Mountains that were reported to me back in 2008 by Joyce Gross (who works for the U.C. Berkeley Natural History Museums and is an avid wildlife photographer). These three trees stood by themselves, with the nearest Palmer’s oaks over 100 miles away to the south in the southern New York Mountains. The next nearest stand of these oaks occurs in the Santa Rosa Mountains, southwest of Palm Desert. Yet, all three stands of oaks share exactly the same species of cynipid wasps. How could this happen? There is no way that these weak-flying wasps could have migrated across the vastness of the Sonoran and Mojave Deserts and colonized these new hosts, so how did they establish themselves on their hosts? The most likely answer came with the discovery of a clone of Palmer’s oaks in the Jurupa Hills, just east of Mira Loma and north of the Santa Ana River. A team of scientists from U.C. Davis and U.C. Riverside, led by Drs. Mitchell Provance and Andrew Sanders, dis- covered a genetically identical clone of these oaks that they estimated to have started growing during the late Pleistocene Era over 13,000 years ago. These researchers think that climatic conditions at the time were more favorable to the growth of Quercus palmeri and that their range was more contiguous, at least in Southern California. With a warming trend developing since the last glacial period, the range of this oak and perhaps other relic populations like singleleaf pinon (Pinus monophylla) have dramatically shrunk, leaving small, disjunct groups of these trees scattered in California. The idea of Palmer’s oaks existing in a more contiguous range is supported by the occurrence of the same species of cynipid wasps now found in these relic groups of trees. Had Palmer’s oaks existed in a broader range, such as from the interior mountains of Southern California across what is now the Mojave Desert to the southeastern Sierra Nevada Mountains, then the occurrence of the same species of cynipid wasps from west to east seems more plausible. Such is the case with the occurrence of the exact same species on blue oak (Q. douglassii) along the Central Valley foothills from Redding to Bakersfield. The small, dependent populations of gall wasp species on Palmer’s oaks have existed generation after generation for thousands of years on a single host tree or small groups of trees. These isolated, relict populations of Palmer’s oaks have now be- come islands in a sea of desert life. While we have managed to travel into space, the grand vastness of the unknown remains nearby in the Mojave and Sonoran deserts.

Deserts remain the “New World” to much we still have to learn about Five new gall species have been discovered me, revealing exciting discoveries the fabric of desert life. on Palmer’s oak since 2009. Palmer’s oak fuzzy gall wasp (Trichoteras burnetti) and with each trip. Lyon’s gall wasp (Heteroecus lyoni) (TOP AND The discovery of new plant galls Ron Russo, 4960 E. 12th Drive, Belling- MIDDLE) were previously known, but the in the Mojave is a reminder of how ham, WA 98226, [email protected] horn gall (BOTTOM) is a new species.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 45 LEARNING TO READ A LANDSCAPE by Phil Van Soelen

eing able to “read” a landscape both in the wild and in one’s own survival often depended upon them. is my personal goal when garden. As a naturalist and gardener, For instance, by observing, our an- gardening with natives. In that has become second nature to cestors were able to recognize drain- practice, this means placing me, but it is a skill anyone can learn. ages where water persists just below Bplants in locations that approximate It also requires some understanding the surface in the dry season. This their native and natural needs. Do- of natural history—of phenomena would have allowed them to spot ing so greatly increases their chances such as sun exposure, landforms, the combination of soil, moisture, of survival over time—and even and drainage patterns. Sometimes and exposure to the sun that favors thriving—without the constant in- at the nursery I talk with people edible roots and bulbs or that might put of water or other resources such who have little idea what the sun provide a life-giving drink of water as fertilizers and pesticides. If a plant does in their garden, or how that in times of need. we’ve placed in our garden thrives changes through the seasons. It’s So how does one develop such without too much attention from also important to know the con- skills in today’s world? Over time I us, then we’ve done our homework tours of your land and how it drains. have taught myself to hone my own well. You probably won’t make wise gar- observational skills for native plant dening decisions without that gardening by “reading” the land- knowledge. scape in the wild. I attempt to un- WHAT IT MEANS TO As a society we have generally derstand the patterns I observe in PONDER lost this skill and knowledge, largely nature and to cultivate the patience because so many people lack any to look deeply by questioning what To read a landscape successfully sort of personal relationship with I see. requires the ability to patiently ob- nature. Acquiring these skills was For instance, why does a par- serve what is happening over time, necessary for our ancestors; their ticular species grow on a ridge as

LEFT: Ferns, manzanitas, dudleyas, and the winter’s flowing water in author’s drainage ditch. These native plants are chosen to function as and to evoke a wild seasonal stream in Sonoma County. They deal with shallow soils, no irrigation, and six months of drought, each in their own way. Photograph by the author. • BELOW: Silver-gray dudleya (Dudleya brittonii), red fuchsia flowering gooseberry (Ribes speciosum), yellow western buttercups (Ranunculus occidentalis), and baby-blue-eyes (Nemophila menzesii) bloom on this seasonally moist bank in the author’s garden. All exist with infrequent to no summer irrigation. Photograph by Saxon Holt.

46 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 opposed to lower down? So Arctostaphylos densiflora)— often in California you will natural hybrids of the com- see a hillside where there is mon manzanita and the oak woodland, then grass- Vine Hill manzanita—on land, and then chaparral. the edge of the ditch, a spe- Through years of experience cies that is tolerant of mois- observing wild landscapes, I ture, good drainage, and know that the oak woodland summer dryness. What I’ve is growing where there is a ended up with is a lovely deeper soil profile with less seasonal creek of manzani- sun exposure, whereas the tas and ferns that is well chaparral is usually growing adapted to the changes of in more rocky soil and has the seasons. a high sun exposure. And Another plant I’ve uti- grasslands are somewhere in lized in my garden is a between. One of the things I dudleya hybrid (Dudleya x enjoy about hiking is con- ‘Frank Reinelt’, also known stantly watching the land- as ‘Anacapa’) from the scape change as you walk Channel Islands. It handles through it, and trying to in- some of same issues in a terpret what is going on. different way. As a succu- This ability to ponder lent, it tolerates our sum- what is happening in nature mer drought fine, but was a skill that was honed plumps up during the win- throughout human history, ter rains. I now have large but today only a compara- drifts of this in much of tively few develop it. In na- my garden. It has created a ture, I find myself encour- Native animals are often seen as pests, but they can function lovely coastal bluff effect aged to ask questions about in a native plant garden much as they do in the wild, with a that needs virtually no sup- what causes whatever it is certain beauty and harmony. plemental irrigation. I’m seeing. Why does a plant grow there? What do the bees or APPLYING THE SKILLS hummingbirds feed upon in the OFFERING CUSTOMERS spring versus in the summer or fall? For example, in my Sebastopol ADVICE Are birds making use of my garden? garden, the previous owners had in- How do the roots from different stalled French drains that channeled At the native plant nursery I co- plants compete with each other? the water into one corner of the own in northern California, custom- How does cold fall down on the garden. This produced an ugly drain- ers often ask us for advice regarding land from above? How does water age ditch, sometimes full of rushing what is the right plant for a particu- flow across the land? Our gardens water in the winter, but bone dry lar site. Over time we have devel- encompass an incredibly complex all summer long. I had to contem- oped a series of questions we ask system of such interactions. plate that as a design challenge. How them in order to guide their choices Ultimately, reading a landscape would I use plants to soften the ditch, (see sidebar). These are the same is about developing a relationship to camouflage it? questions I almost unconsciously ask with nature, a process I find satisfy- My solution was to create a fern- myself when placing plants in my ing and that eventually becomes sec- lined seasonal stream by planting own garden. For instance, custom- ond nature. Once one begins to ac- California polypody (Polypodium ers may tell us their garden is prima- quire this observational knowledge, californicum)—a native fern one sees rily in shade. Most plant labels only it proves extremely helpful in guid- in nature along streams and canyon differentiate between shade, partial ing one’s gardening choices about walls, and that goes summer dor- shade or sun, or full sun. At first what plants to select and where to mant—on both sides of the drain- glance it would seem a fairly easy place them in the home garden. The age ditch. When the ditch dries up decision to place shade-loving plants same process can then be used to each summer, the fern tolerates that in a shaded part of the garden. But to evaluate how well they respond, and perfectly. I also planted several man- truly read a garden landscape in or- to modify future gardening choices. zanitas (Arctostaphylos manzanita x der to know what to plant and where

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 47 SOME QUESTIONS TO GUIDE THE NATIVE PLANT GARDENER 1. Where in the county is your garden (coastal, inland, in the hills, valley bottom)? 2. What is its sun exposure? (Are there trees or structures blocking the sun? Does it have open or dappled sunlight? What are its qualities of shade—bright, deep, etc.—and what is creating the shade?) 3. Are you going to irrigate, and how much, especially in the sum- mer? How much available moisture already exists in places? 4. Are there deer or gophers in the area? (These are the two major pest animals in our county.) 5. What is the quality of the soil? (What color is it, i.e., dark choco- late brown, light beige, gray? You want to try and get some sense of whether it contains primarily clay, sand, or a mixture of the two, so you can help gardeners make intelligent plant choices based on drainage and soil types.) 6. Do you want primarily natives, or natives and non-natives that are drought tolerant? 7. Are you comfortable with silver-gray foliage, or do you prefer green and dark-green leaves? (This will have a major impact in terms of the plants I recommend.) 8. Do you like things very tidy and formal, or more naturalistic and informal?

to plant it, we need considerably more information than that. There are different degrees of shade, such as bright, high-canopy deciduous shade, versus deep ever- green forest shade. We need to know what type of shade exists in that particular spot in the garden. There are a number of other factors that also should be taken into account before making any planting deci- sions, such as the amount of avail- able moisture, the type of soil and available nutrients, the proximity to est garden spot where water col- large trees or shrubs, how much lects naturally, and determine if it wind the site gets, and the angle and also provides other requirements amount of sun exposure in different of these plants, namely bright shade Many of the same forces—substrate, seasons. yet little or no root competition nutrient flows, and exposure to sun—that So, for example, if a customer from other large plants or trees. operate on a larger scale in the home says she would like to plant trilli- Knowing what a plant requires for garden, also stratify and segregate vege- tation patterns on the micro level, in this ums or lilies in her garden—plants optimum health and being able to case lichens on a rock at the Pinnacles that want moisture and bright find the optimal location for it in National Monument. shade—we need to locate the low- the garden are key to creating a

48 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 Sun exposure, drainage patterns, existing vegetation, and soils all combine to make a rich and pleasing mosaic on Mount Diablo. Nature is the best instructor in planning our home gardens. beautiful and successful native plant a dormant state and leaf out just as garden. But for me, perhaps the most garden. the angle of the sun raises enough to compelling is being able to recreate Observing one’s own garden provide adequate light for photosyn- outside my home the appealing inti- keenly throughout the seasons also thesis. This allows planting in areas macy I sometimes experience when reveals valuable information. As an darkened by the low angle of the sun I’m in the wild, the feeling that “this example, many shade tolerant plants in the winter—more so than many spot feels so right.” The ability to are deciduous (they lose their leaves evergreen plants would tolerate—yet read a landscape makes this possible. seasonally) or herbaceous (they go it still allows them to receive bright dormant back to their roots). They shade or sun in summer. Phil Van Soelen, 7560 Brittain Avenue, survive the darkest days of winter in There are many reasons people Sebastopol, CA 95472, [email protected]

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 49 DEVELOPING A SUSTAINABLE MIX FOR SEED GERMINATION USING LOCAL MATERIALS by Jackie Bergquist Salas, Michelle Laskowski, Brianna Schaefer, and Betty Young

he Golden Gate National Peat moss has been a major compo- blend produced within the park in Parks Conservancy’s Native nent of both media. Unfortunately, 2003. With careful control of the Plant Nurseries are charged peat moss—one of the most com- compost quality, transplanted and each year with the produc- mon amendments in germination mature native plants have performed tionT of 150,000-200,000 native and growing media—is harvested on very well in this compost-based mix. plants grown from watershed-spe- a scale much faster than peat bogs However, so far we have been un- cific seed, which are used in restora- replenish themselves and as such, is successful at formulating a seedling tion projects throughout this na- not very renewable (Priesnitz 2007). germination medium. tional park. The Nurseries’ goal is to Additionally, peat moss is harvested For the past few years, with fi- produce plants in the most ecologi- and shipped from Canada. Coconut nancial support from the Presidio cally sustainable manner possible. fiber (coir), another substitute for Trust, we have been trying to de- This commitment includes consid- peat moss, is shipped thousands of velop an alternative seed germina- ering the ecological footprint of each miles to reach our nurseries in the tion medium with a rate of germina- product used in plant production. San Francisco Bay Area. Shipping tion similar to that of the commer- The main products used during either peat or coir uses a large cial peat moss-based medium, while the production process are seed ger- amount of fossil fuel. using local sustainable amendments. mination media, used in 16" x 16" x Due to these ecological and en- In 2009, after a review of experi- 2" flats, and potting media for grow- vironmental concerns, the regular mental literature, we chose compost, ing seedlings that have been trans- potting medium used at the Nurser- partially boiled rice hulls (a by-prod- planted into their final containers. ies was converted to a compost-based uct of California’s agriculture indus-

TABLE 1. GERMINATION TRIAL MEDIA RECIPES

Trial Medium #1 Trial Medium #2 Trial Medium #3 Ingredients Percentage Amounts Percentage Amounts Percentage Amounts Sifted Compost 33.30% 12 L 33.30% 12 L 25% 9 L Boiled Rice Hulls 0% 0 L 33.30% 12 L 25% 9 L Earthworm Castings 33.30% 12 L 0% 0 L 25% 9 L Perlite, fine grade 33.30% 12 L 33.30% 12 L 25% 9 L 36 Liters 36 L 36 L Fertilizers added to all trial mixtures: Oyster Shell 6.00% 23g Soybean Meal 11.70% 45g Iron Sulfate 2.30% 9g Gypsum 4.40% 17g Sustane™ 4-6-4 75.60% 291g Total 100% 385g

Trial Medium #4 – Control. Commercial medium (Sungro Horticulture Sunshine Formula 5), includes Canadian sphagnum peat moss, fine perlite, gypsum, dolomitic lime, and a low fertilizer charge. Results of our trial media were measured compared to this mix.

50 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 try), and earthworm castings as local and sustainable materials that could potentially provide the basic properties of a successful growing medium (Hidalgo et al. 2009, Hoitink et al. 1997, Evans R.Y. 2006, Merhaut 2006, Evans M. 2008, Merca 2009). Since performance of rice hulls in germination media was unknown, perlite was added as a porous/structural component to maintain sufficient aeration in the mix. Media mixtures were prepared to test the amendments in different combinations (see Table 1, Germina- tion Trial Media Recipes). Samples of the media were then sent to an independent laboratory for analysis of their physical and chemical prop- erties. Flats were sown with three species of seed chosen for their vary- to the other media evaluated ing characteristics: California aster (see Table 2, Results). Al- (Symphyotrichum chilensis, also though the control medium known as Aster chilensis), blue wild performed best in terms of rye (Elymus glaucus), and sticky percentages, statistical data re- monkeyflower (Mimulus aurantia- vealed no significant differ- cus). These seeds were sown in ran- ence between the germination dom on standardized and grid pat- rates of Mix #1, #3, and the terned germination flats that re- control medium. However, the ceived intermittent mist set at a tem- germination rates in Mix #1 perature of 65�F (18.33�C). Flats and Mix #3 were only around were monitored to note the date of 75% of the germination rate germination, potential mortality, of the control peat-based mix- and/or the date seedlings were large ture. enough to transplant. This data was In regard to physical prop- collected three times a week for erties, Mix #1 was dense and seven weeks. became water logged, while The results of the 2009 trial show Mix #3 with compost and rice that the control peat-based mixture hulls had better drainage, but produced the highest germination did not supply nutrients rate, fewest weeds (germinating from quickly enough to the seed- weed seeds in the various amend- lings, delaying growth. The ments), lowest overall mortality earthworm castings in Mix #1 rates, and largest fresh and dry and #3 contained weed seeds weight of roots and shoots. Trial that had to be pulled. While Mix #1, which was equal parts earth- the rice hulls break down worm castings, compost, and per- slowly, holding their shape in lite, and Mix #3 comprised of these ingredients plus boiled rice hulls TOP: Developing a sustainable mix for seed germination using local materials. Left to right: seemed to perform nearly equally Mix #1 (earthworm castings, perlite and compost), Mix #3 (earthworm castings, perlite, well. Mix #2, equal parts rice hulls, compost, and boiled rice hulls), Mix #2 (compost, boiled rice hulls and perlite). All photographs by Jackie Bergquist Salas. • MIDDLE: Flats of each mix planted with three compost, and perlite, lacked nutri- native species, including sticky monkeyflower, blue wild rye, and California aster, six ent and water holding capacity, and weeks after sowing. Clockwise from top left: Mix #3, Mix #2, Mix #1, and Control. • did not perform well in comparison BOTTOM: Preferred experimental mix: Mix #1 after seven weeks.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 51 TABLE 2. RESULTS OF EXPERIMENT BY SPECIES AND MIX

Mortality Root Root Shoot Shoot Root: Germination Number Averages % of Fresh Dry Fresh Dry Shoot % Harvested seedlings Weight Weight Weight Weight Ratio Mimulus Mix # 1 53.54% 2.57% 16.00 0.46 0.16 3.83 0.53 0.30 Mix # 2 25.25% 42.03% 4.33 0.01 0.10 0.03 0.04 n/a Mix # 3 35.35% 27.37% 9.67 0.53 0.11 1.96 0.31 n/a Control 52.53% 3.67% 17.00 0.59 0.30 6.55 0.91 0.33 Aster Mix # 1 23.23% 0.00% 7.67 2.52 0.43 8.68 1.02 0.54 Mix # 2 35.35% 25.47% 8.67 0.81 0.18 1.48 0.29 0.48 Mix # 3 41.41% 18.07% 11.00 3.55 0.56 8.02 0.78 0.71 Control 52.53% 0.00% 14.67 15.54 2.03 29.46 4.05 0.53 Elymus Mix # 1 60.61% 6.20% 18.67 15.86 2.88 30.40 5.63 0.53 Mix # 2 57.58% 7.70% 17.67 3.92 0.84 5.03 1.06 0.85 Mix # 3 63.64% 3.10% 20.33 14.68 3.08 20.62 3.95 0.83 Control 71.72% 0.00% 23.67 91.65 13.29 48.53 9.49 1.38 the medium, the absence of earth- hearing from other nurseries or Merca, D. 2009. Personal communica- worm castings in Mix #2 created a growers about how they have made tion. California Rice Commission, nutrient poor mixture for develop- their media more sustainable. Sacramento, CA. ing seedlings. Merhaut, D.J. 2006. Substrates and their use. California Ornamental Research The results of this trial show that REFERENCES with minor adjustments, compost, Federation News 10(3): 1-2, 11. Nelson, D.V. 2003. Chapter 6 in Green- earthworm castings, and partially Evans, R.Y. 2006. Compost as a pot- house Operation and Management. boiled rice hulls can perform well ting mix amendment: Too much of a Pearson Education, Inc., Upper when combined to make a germina- good thing? California Ornamental Saddle River, NJ. tion medium. Our next experiment Research Federation News 10(3): 12. Priesnitz, W. 2007. Does peat moss will be to modify Mix #3 with one Evans, M. January 2008. Rice hulls 101, have a place in the ecological gar- part compost, one part earthworm Grower Talks. Ball Publishing, Chi- den? Natural Living Magazine. Ac- cago, IL. castings, and two parts boiled rice cessed January 26, 2010. http://www. Hidalgo P., M. Sindoni, F. Matta, and hulls. This increase in rice hulls naturallifemagazine.com/0712/ D.H. Nagel. 1997. Earthworm cast- asknlpeat.html. should improve drainage, and the ings increase germination rate and exclusion of perlite will make it more seedling development of cucumber. Jackie Bergquist Salas, Children’s Fairy- sustainable. We will sift the amend- Mississippi Agricultural and Forestry ments through 1/4" hardware cloth Experiment Station, MSU Cares Re- land, 699 Bellevue Avenue, Oakland, CA to provide a more uniform mix and search Report 22, No. 6. Accessed 94610, [email protected]; adjust the fertilizers slightly for a June 18, 2009. http://msucares.com/ Michele Laskowski, Golden Gate National quicker release. pubs/researchreports/rr22-6.htm. Parks Conservancy, 1216 Ralston Avenue, We demonstrated that it is pos- Hiotink H.A.J., M.A. Rose, and R.A. Presidio of SF, San Francisco, CA 94123, sible to make local sustainable re- Zondag. 1997. Properties of materi- [email protected]; als available for formulation of high- sources compare favorably with con- Brianna Schaefer, Golden Gate National quality container media. Ohio State ventional non-sustainable products. Parks Conservancy, 1216 Ralston Avenue, University, Ohioline Bulletin, Orna- Presidio of SF, San Francisco, CA 94123, With some simple modification to mental Plants Annual Reports and Re- the recipes, we are confident that search Reviews Special Circular 154. [email protected]; Betty we will soon be able to produce a Accessed June 18, 2009. http:// Young, Golden Gate National Parks Con- sustainable germination medium for ohioline.osu.edu/sc154/sc154_14. servancy, 201 Fort Mason, San Francisco, our Nurseries. We would appreciate html. CA 94123, [email protected]

52 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 Roger in 2011 at The Cedars with his beloved dogs, Serpentino (gray), Hortus (black), and Daisy (tan). The Cedars has been a focus of Roger’s life since 1981 when he first entered its canyons. Photograph by David McCrory. NEW CNPS FELLOW: ROGER RAICHE by Phyllis Faber

oger Raiche has made unusu- veloped an extraordinary serpentine The Cedars. In sum, Roger’s life has ally extensive contributions garden. been devoted to California’s flora to California botany in four As a horticulturalist, Roger made and to sharing his love for this flora. distinct arenas. As a foremost extensive collections of native seed, Roger worked for 23 years at the RCalifornia botanist, he has gener- always with precise collection data, U.C. Botanical Garden (UCBG) in ously shared his extensive knowl- collected numerous plant variants, Berkeley in charge of the 15-acre edge with many audiences, includ- and developed several interesting California Collection. He was the ing numerous members of CNPS, cultivars for gardens. And as a land primary collector of that section for and has discovered several new na- protector of the best sort, he pur- all those years and brought in thou- tive species that have subsequently chased a core area of “The Cedars,” sands of wild plants during that time been named for him. As a gardener an extraordinary serpentine barren with precise collection data. He was par excellence, for years he main- in Sonoma County in order to pro- also primary collector for the Gar- tained the California Native section tect it. He has been host to many dens’ Index Seminum or seed list of the University of California Bo- CNPS members and to fascinating that is put out every two years and tanical Garden at Berkeley, and de- and delightful chapter field trips to goes to over 500 botanical institu-

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 53 tions around the world (for a total mented on numerous plant entries has focused on plants growing on of ten lists during his tenure at the in that volume. that substrate. Protecting and un- Garden). Although most of his Perhaps Roger’s most impressive derstanding an area in Sonoma collecting was done for UCBG, he talent regarding native plants was County called The Cedars became conducted most of it on his own his ability to find them, especially part of his obsession, and he became time. rare ones in the wild, as well as a vigorous advocate for its conser- Roger wrote a few articles over finding new ones. Three new taxa vation since 1983. He has led many the years pertaining to natives, in- were named in his honor: Cedars field trips for CNPS to this site, as cluding an article in Pacific Horti- fairy-lantern (Calochortus raichei), well as giving numerous slide lec- culture in 1991 describing many of Raiche’s red ribbons (Clarkia con- tures over the years pertaining to his cultivars. He also wrote a col- cinna ssp. raichei), and Raiche’s man- the native flora growing on this huge umn called the “California Corner” zanita (Arctostaphylos stanfordiana serpentine area which has been for a monthly internal UCBG news- ssp. raichei). Roger was also very eroded into many deep narrow can- letter called the UC Bee, where he talented at finding variants that he yons. At least seven distinct plants wrote about various things of inter- thought might make interesting cul- are found only at The Cedars. est happening in the California area. tivars for horticulture, including In 1999, Roger and his partner, Roger was also on the Jepson Manual around 20 that he named. Roger’s David McCrory, purchased 520 acres Horticultural Council, which helped introductions of exceptional native in the heart of The Cedars. This prepare the horticultural entries for plant cultivars frequently showed up allowed them to restore damaged The Jepson Manual (1993) that War- for sale at the CNPS East Bay Chap- areas, to continue with field trips, ren Roberts chaired. Roger com- ter’s plant sales before they had re- and to facilitate scientific studies. In ceived wider circulation. This cer- 2011, in a collaborative effort with Roger with his business partner, David tainly had the effect of supporting Save The Redwoods League, the State McCrory, at The Cedars. Photograph by the chapter by making the sales more Coastal Conservancy, and the Gor- Phil Van Soelen. interesting. don and Betty Moore Foundation, Several native genera have been 500 acres of this important core area of special interest to Roger, espe- were purchased and transferred to cially Arctostaphylos, Ceanothus, and BLM (Bureau of Land Management) Streptanthus. He made many hun- to add to its Area of Critical Envi- dreds of herbarium specimens of ronmental Concern at The Cedars. those and other taxa over the years. Roger also helped the Sonoma Land This work helped to identify many Trust (SLT) develop its “Conserva- new sites for existing and possibly tion Plan for The Cedars Area,” some new taxa. In recent years, which will guide future conserva- Roger collaborated with Dr. James tion efforts of this region. Reveal of Cornell to publish two Warren Roberts, Superintendent new taxa that occur at The Cedars, of the U.C. Davis Arboretum (now The Cedars buckwheat (Eriogonum retired), and former president of the cedrorum) and The Cedars ocean- California Horticultural Society, de- spray (Holodiscus dumosus var. cedro- scribed Roger as follows: rus). Monocots also really interested Roger Raiche fell in love with the him, and he was a major contributor plants of California literally at first to the CNPS book, Wild Lilies, Irises, sight. Driving across the Sierra en and Grasses: Gardening with Califor- route from his home town, New- nia Monocots, finally published in port, Rhode Island, to his new life 2003 after 30 years of a working in California, he was struck by the study group. Roger did the primary beauty of our native manzanitas. writing for Brodiaea, Dichelostemma, He never got over it. His enthusi- Trietleia, Fritillaria, and all the asm for the flora of his adopted Grasses, Sedges and Rushes—a ma- home state has grown over the jor component of the book. years, and for decades he has been One of Roger’s major obsessions one of California’s greatest advo- has been the serpentine flora of Cali- cates for the beauty and garden- fornia, and much of his collecting worthiness of our native plants.

54 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 A vintage photograph of Roger back in 1987, standing in a chaparral (on Cow Mt. east of Ukiah) of the Raiche manzanita (Arctostaphylos stanfordiana ssp. raichei). The species was named in his honor that year by Walter Knight of the California Academy of Sciences. Photograph by Roger Raiche.

Phil Van Soelen, Cal Flora Nurs- ley Botanic Garden, the California ery co-owner in Santa Rosa, and a native area was transformed from a past president of the Milo Baker nice, rather unexceptional garden Two of the 20 cultivars that Roger has propagated and named. TOP: Chapter of CNPS, made the follow- of natives to a very dynamic artistic ‘Joyce Rose’, a deep rose selection ing observation of Roger: and scientific creation. Roger has a of the coast flowering currant, very rare blend of exceptional ar- Ribes sanguineum var. glutino- Roger really is a towering figure in tistic abilities as well as acute sci- sum, which he named in honor horticulture in general, and Cali- entific perception and memory. He of his mother, Joyce Rose Raiche. MIDDLE: The magnificent burgun- fornia native plant horticulture in has always been extremely gener- dy leaf stream orchid, Epipactis particular. I believe he took the ous with his time and energy lead- gigantea f. rubrifolia ‘Serpentine mantle directly from Wayne ing CNPS field trips. My business Night’. Both photographs by Phil Van Roderick. A state CNPS survey partner Sherrie Althouse said that Soelen. • BOTTOM: Roger has demonstrated showed that, among all respondents Roger led the best field trips she an uncanny ability to find rare and new native plants in the wild. One of three new who answered it, members’ inter- has ever been on, and from my taxa named in his honor is The Cedars est in’gardening placed at the top experience I would agree. His trips fairy-lantern (Calochortus raichei). Photo- of the list of topics. Certainly gar- to the Cedars are legendary. graph by Roger Raiche. dening with native plants was first developed and showcased to Roger Raiche has perhaps been preciating our native flora, making it the public in botanic gardens, and one of the finest and most effective known to others, and protecting it. Roger’s contributions in that field native plant advocates of this gen- are great. There really is no one eration, and California has been for- Phyllis M. Faber, 212 Del Casa Drive, else of his caliber and accomplish- tunate to have his skills, energy, and Mill Valley, CA 94941, pmfaber@comcast. ments. While he was at the Berke- devotion in understanding and ap- net

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 55 SCOTT FLEMING: 1923–2011 by John Danielsen

cott, with his wife Jenny, was chapter and state CNPS board meet- garden infrastructure, Jenny took the among the founding mem- ings, or conservation strategy and lead in acquiring the plants. Through bers of CNPS in 1965, and planning sessions held in Berkeley this teamwork they created a home became a Fellow of the Cali- or hosted at their home. garden that remains one of the sig- Sfornia Native Plant Society in 1985 Scott loved to travel on field trips nature native plant gardens in the (see the April 1985 issue of Fre- to the far corners of the state, and Bay Area. montia). Scott was active in CNPS we shared this passion with Scott The effort to create their garden, at the state level as treasurer and and Jenny. Scott also enjoyed shar- plant by plant and rock by rock over long-time legal advisor, where he ing meals and conversation with many years, also reflected the dedi- brought creative friends, and he was an expert chef cated plant conservation ethic they thinking to solve with his famous in-house smoke both shared and engendered within the Society’s early oven. When Scott decided to step CNPS. And as we remember our financial growing down as state CNPS treasurer, he dear friends, one legacy of their love pains, as well as asked me to take on the treasurer lives on in the beautiful garden they negotiating the position. He was always ready to generously shared with all. contract for CNPS provide good and insightful help as to produce its first CNPS nearly doubled in size over John Danielsen, 10 Kerr Avenue, Kensing- plant poster. the next few years. He was equipped ton, CA 94707, [email protected] A lawyer by with that unique quality to be able training, Scott to listen carefully to what the prob- Scott and Jenny in their Bay Area native used his skills to lem of the moment appeared to be, plant garden. put together, with rephrase it in a way that helped oth- others, the Kaiser- ers understand, and then suggest Scott Fleming. Permanente creative solutions. Health Program, I often think of a raft trip we and helped to found the Planning took down the Colorado River in and Conservation League, where he the early 1990s. Scott and I hosted worked tirelessly promoting conser- about 25 people on a two-week rub- vation activities in California. Scott ber raft trip through most of the was an avid white water kayaker who white water available to boaters. We also enjoyed hiking and camping. traveled in small rafts, each holding Scott and Jenny are survived by about five persons. At one point on daughters India and Hilari as well as the river we were faced with mon- three granddaughters. strous waves going over Lava Falls. I first met Scott on one of our Observing these from the banks of chapter’s weekend hikes, hoping to the river, one person commented, discover the wonders of the Bay “Holy smokes, I am not going over Area’s native flora. We also discov- those,” to which Scott replied, ered that we worked for the same “Watch me, I’m going over them in company, Kaiser-Permanente of a kayak!” Well, we all watched Scott Oakland, and that our spouses were make it, so the rest of us followed, good friends. Scott and Jenny were a inspired by his example. Scott was wonderful and inspiring team who capable, fearless, and expert on these warmly welcomed my wife Charli trips, which were wonderful quali- and me into their home and family. ties to have in a friend and colleague. They continued over many de- Scott and Jenny loved to garden, cades to provide this hospitality to although truth be told, Scott spent other CNPS members from all over some of the garden time doing kayak the state who were in need of a good rollovers in the family pool. While sleepover while attending the many Scott was the prime builder of the

56 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 RALPH MILTON INGOLS: 1911–2011 by Leah Price Hawks

e have lost one of the ing, and journalism. During this finest conservationists time he received his master’s degree that one could ever and counseling certification at San have known. Centenar- Francisco State University. Ralph ianW Ralph Milton Ingols passed away coached numerous sports including in Napa, California on July 11, 2011. football, basketball, baseball, wres- A product of Weed Patch in Kern tling, and boxing. He also gave les- County, California, Ralph was the sons to the school ski club at his youngest of seven siblings. He and cabin in the Sierra Nevada. Ralph his wife Evelyne were parents of the and Evelyne were active in the Napa “Three J’s,” Janet, Jim, and Judy Valley Chapter of the California Na- Ingols, to whom they were devoted. tive Plant Society, and for many years Their children joined them in a life- used their backyard to propagate time of outdoor activities, which native perennials for the chapter’s cemented their interest in the envi- plant sales. After they both retired Ralph Ingols in the mid-1950s at St. Helena ronment as well as in education. in 1972, the couple spent many years High School, where he taught for over 30 As a toddler, Ralph was often traveling throughout California, years. put on a blanket among the wild- studying and collecting its native flowers and grasses in Weed Patch. flora and gaining valuable knowl- servationist friends, the Napa Valley His ground-eye view of the land- edge and background in plant iden- Chapter of CNPS, and the Skyline scape left a lasting impression on tification and propagation. Citizens Association, they founded him and was his earliest instructor During this period, the Ingols the three-acre Martha Walker Na- in the natural world. At a young age, and their friends first began discuss- tive Habitat Garden, located at Sky- Ralph knew he wanted to become ing the creation of a public garden in line Wilderness Park, and became an educator, and later on he did. the Napa Valley where visitors could its first curators. Ralph and Evelyne He received his B.A. and teach- learn about and enjoy California’s were the driving force behind the ing credential at U.C. Berkeley and flora in protected surroundings. The new garden in its beginning years, then began a 35-year career—most idea of a garden that could be used and Ralph continued as curator of which was spent at St. Helena as an outdoor classroom particularly emeritus after Evelyne’s death in High School in the Napa Valley— appealed to the Ingols, since both 2000. teaching California and world his- were career educators. Those close to Ralph knew that tory, world literature, public speak- By 1985, with the help of con- he always had a broad vision for the

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 57 also finished five other books dur- ing this time, two with coauthors, including one about St. Helena High School, and another on Skyline Wil- derness Park. Proceeds from the sale of his books go to support CNPS and the Martha Walker Garden. Just a few years ago, Ralph re- ceived the Earl Thollander Conser- vation Award, presented annually by the Napa Group of the Sierra Club to an individual who has achieved excellence in conservation. Perhaps one of Ralph’s most im- pressive accomplishments was his having the Martha Walker Garden named a World Peace Garden in 2002, the first in the United States ABOVE: The Ingols, both teachers, believed strongly in using the Martha Walker Gar- to receive that honor! (Each garden den as an outdoor classroom. Ralph, one in the England-based World Peace of the garden’s science docents, had a knack Garden Project serves as a retreat for sharing with students his love of plants for peaceful reflection on the beauty and animals. • LEFT: Ralph with his wife, of nature and the ideals of living in Evelyne, in the Sierra Nevada in the mid- 1990s. The two were the driving force harmony with others.) In 2010 Ralph behind the creation of the Martha Walker attended the celebration of the Native Habitat Garden in Napa County. Garden’s 25th anniversary, and up until a few years ago, he also at- In his later years, Ralph contin- tended as many class reunions of his ued to make every effort to try new high school students as he could. things at an age when most people Ralph was planning for the fu- consider their life’s work finished. ture up until the day before he passed He wrote, planned, and organized away, still making arrangements for the community to accomplish wor- the Garden to benefit in yet more thy goals, and in the process con- ways through plantings, work par- nected with others in every walk of ties, and proceeds from his books. life. “When people go into the Garden, Ralph always spoke warmly of I want them to see butterflies every- Garden. “Future generations can others he respected, whether com- where,” he said. “I want to plant enjoy what we prepare for them to- munity leaders, former students, vol- Dutchman’s pipevine for them.” day!” was his motto. Thanks largely unteers, or youth, and never missed His friends in the Napa Valley to his efforts, the Napa Valley Chap- a chance to encourage others. In Chapter are now working to make ter established a science docent pro- one way or another, Ralph was a that dream come true. Additional gram there. Today that program en- mentor to everyone he met. He sim- milkweed to attract monarch but- sures that young students who visit ply felt that it was within the capac- terflies and Dutchman’s pipevine the Garden learn about its plants ity of each person to succeed, and to attract the pipevine butterfly are and wildlife. he was capable of convincing gen- just two of the species being dug in Ralph spoke of a time, a hun- erations of students and friends that now, and a steering committee is dred years from now, when this this was so. planning a Ralph Ingols Butterfly generation’s grandchildren would Over the past ten years Ralph Trail to be completed in 2012 to also walk the paths of the Martha published two books for his family: honor Ralph’s dedication and vision Walker Garden under his favorite The Ingols Cabin, stories of fam- for the Garden. trees. He willingly gave whatever he ily and student adventures in the could in energy and resources to the Sierra, and Why I Am Who I Am, a Leah Price Hawks, 1236 Second Avenue, community, and in particular to the biographical sketch of his life in Cal- Napa, CA 94558, leah7hawks@yahoo. Garden. This was his legacy. ifornia. He was a prolific writer, and com

58 FREMONTIA VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 LETTERS TO THE EDITOR

WETLANDS: MISPLACED there is so precious little natural wet- as it has been for decades—is simply land habitat left—probably well less to save as much suitable wetland- EMPHASIS than 5%, and even that is severely frag- supporting land as possible (actual mented—I believe we no longer even and potential), enhance it as needed, Having studied the seasonal wet- have the same underlying ecosystem funded, and practical (including plant lands of Sonoma County’s Santa Rosa within which to manage the remnants, translocation), and lock it away. Do Plain (SRP) for the last 27 years (con- and that continued habitat construc- we need to improve our preserve man- sulting to developers, city, county, and tion/restoration with rare plant trans- agement? Absolutely. Do we need more state agencies, preserve proponents, location will likely be important parts research? Always. I’m not sure if these nonprofits and landowners, and as a of any hoped for recovery. are desperate times calling for desper- congressional appointee to the Sonoma Wetland (and hence native pool ate measures just yet, but in some re- County Vernal Pool Task Force, 1988- flora) conservation on the SRP has spects it feels very close. 1992), I am glad to see the attention already come a long way since the Charlie Patterson being paid to this unique area (see 1980s, thanks largely to people like Plant Ecologist, Lafayette, CA Fremontia Vol. 37, No. 4/Vol. 38, No. Betty Guggolz and Nancy Harrison 1, pp. 40-43). However, I feel com- (both had been active members of pelled to comment. CNPS) and others, plus a more active First, with regard to history, it NEED FOR SCIENCE-BASED role by the agencies, specifically re- should be noted that well before 1970, CONSERVATION quiring compensatory mitigation. The this region’s broad valley of seasonal results to date, while far from perfect, STRATEGIES: THE AUTHORS wetlands, perennial grasslands, oak include the long-term preservation of REPLY savanna, and riparian forests had more than 1,500 acres on more than already been thoroughly (80-90%) 50 preserves. We appreciate Mr. Charlie chopped up, chopped down, con- Finally, while I encourage all sci- Patterson’s perspective in his response verted to prunes, pears, grapes, dair- entific efforts on the SRP, I also be- to our article on the vernal pools of the ies, mustard and annual grassland, lieve that today’s main challenge— Santa Rosa Plain in Sonoma County. and dewatered by roads, ditches, berms, buried drains, and engineered channels that flow straight to the La- guna de Santa Rosa. Up until the early 1980s, federal law allowed for the “minor fill” of up to one full acre of wetland under the “nationwide per- mit program,” often with little or no significant mitigation. Then, with probably less than 5% of the region’s wetlands left, intense efforts were made over the last several decades to reduce, modify, and/or miti- gate modern development’s wetland impacts. For 25 years now, the U.S. Army Corps of Engineers has required at least 1:1 replacement of wetland functions, values, and acreage (essen- tially dictating new wetland construc- tion). In addition, the U.S. Fish and Wildlife Service currently requires the provision of acreage or credits sanc- tioned as “Establishment” habitat, i.e., new physical habitat constructed and seeded with listed species. The region’s initial efforts toward rare plant translocation (e.g., Alton Lane) were conducted at the direction of, and in fact as a permit requirement of the California Department of Fish and Game. Hence, on the SRP, where

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA 59 land as possible. But we regard sci- ence as the most effective means to determine what is “suitable.” Scien- tists estimate that only 5-10% of California’s original vernal pool habi- tat remains, and a great deal of recent conservation efforts have been chan- neled into mitigation, while destruc- tion of natural habitat continues. But if man-made wetlands fail to provide sustainable conditions for translocated populations, then we are faced with the unfortunate reality that vernal pool mitigation may be a slow and expen- sive method for extirpating popula- tions, and eventually the species that we aim to protect. Mr. Patterson, who has personally been involved in numerous mitigation efforts, admits that the results of re- cent conservation efforts may be “far from perfect.” But just how far? Are While the plants of the Santa Rosa ments our forward-looking perspec- they too far to be an effective conser- Plain are threatened by human im- tive on vernal pool conservation on vation strategy? Hopefully not, but we pacts in many ways, they are also the Santa Rosa Plain. Indeed, habitat argue that mitigation decisions—and deeply valued by a large number of loss in the area began well before the their consequences—should be more passionate local constituents that are early 1970s, yet the rapid urban ex- critically monitored, and more heavily eager to contribute to the conserva- pansion during 1970–1990 unequivo- researched. Vetted protocols, appro- tion of the flora. We are aware of Mr. cally pushed the threats facing native priate controls, adequate replication, Patterson’s experience with vernal pool habitats in this region to a new level of and rigorous data analysis can facili- mitigation efforts on the Plain, appre- urgency. As discussed in our original tate the objective evaluation of the de- ciate his genuine interest and concern article, the accelerating loss of vernal gree to which mitigated vernal pools for vernal pool communities, and wel- pool habitat intersected with endan- are supporting sustainable vernal pool come his outlook on our article. How- gered species regulations during this populations and communities. Fur- ever, we respectfully disagree that our time to prompt the regulation of ver- thermore, detailed studies of remnant emphasis on science-based conserva- nal pool mitigation projects by CDFG populations are essential for determin- tion and community involvement on and other agencies. In the meantime, a ing the important biological processes the Santa Rosa Plain is misplaced, and great deal of mitigation, translocation, influencing population growth and we are eager to reiterate our position and habitat loss occurred before the persistence in the fragmented context that scientific research is an essential basic ecological requirements of ver- that now characterizes the vast major- part of ensuring the long-term sur- nal pool species were understood. ity of historic vernal pool habitat. vival of vernal pool habitat. We fully agree with Mr. Patterson The problem, of course, is that The additional history provided in that our main challenge is to save as thoughtful science and careful moni- Mr. Patterson’s letter nicely comple- much suitable wetland-supporting toring takes time, and often must oper- ate at a slower pace than the pressures of development and the rates of habitat loss. Fortunately, there are conserva- tion scientists willing to take that time Telos Rare Bulbs in the Santa Rosa Plain. Hopefully, the The most complete offering of bulbs native to the practitioners of vernal pool mitigation and conservation will facilitate that re- western USA available anywhere, our stock is search by sharing any monitoring data, propagated at the nursery, with seed and plants releasing historical records, and facili- from legitimate sources only. tating site access, and will be eager to develop strategies that incorporate the Telos Rare Bulbs insights from these projects. P.O. Box 1067, Ferndale, CA 95536 Nancy C. Emery, Purdue University www.telosrarebulbs.com Michelle Jensen, Ecologist, Pepperwood Preserve

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CONTRIBUTORS (continued from back cover) Patrick Kobernus, biologist, supervised habitat management and endangered MATERIALS FOR species monitoring on San Bruno Mountain from 1995–2007. He is currently a PUBLICATION consultant for Coast Ridge Ecology, and a member of the Yerba Buena Chapter of Members and others are invited CNPS. to submit material for publica- tion in Fremontia. Instructions Michele Laskowski is seed ecologist for the Presidio Native Plant Nursery in the for contributors can be found Golden Gate National Parks. on the CNPS website, www.cnps. Ron Russo is the retired chief naturalist of the East Bay Regional Park District org, or can be requested from where he worked for 37 years, and the author of Field Guide to Plant Galls of Fremontia Editor Bob Hass at California and Other Western States. [email protected].

Brianna Schaefer is manager of the Presidio Native Plant Nursery in the Golden Gate National Parks. Fremontia Editorial Advisory Board Hugh Safford is regional ecologist for the U.S. Forest Service’s Pacific Southwest Susan D’Alcamo, Ellen Dean, Region, and a research associate with the Department of Environmental Science and Policy, University of California, Davis. Phyllis M. Faber, Holly Forbes, Brett Hall, Tara Hansen, Todd Phil Van Soelen is co-owner of California Flora Nursery, a past president of the Keeler-Wolf, David Keil, Pam Milo Baker Chapter (Sonoma County) of CNPS, and teaches classes at Santa Rosa Muick, Bart O’Brien, Roger Junior College on gardening with California native plants. Raiche, John Sawyer, Teresa Betty Young is director of the six native plant nurseries located throughout the Sholars, Greg Suba, Dick Turner, 80,000 acres of the Golden Gate National Parks in Marin, San Francisco, and San Mike Vasey, Carol Witham Mateo Counties.

VOLUME 38:4/39:1, OCTOBER 2010/JANUARY 2011 FREMONTIA California Native Plant Society Nonprofit Org. 2707 K Street, Suite 1 Sacramento, CA 95816-5113 U.S. Postage PAID Address Service Requested A.M.S.

CONTRIBUTORS FROM THE EDITOR

Earl B. Alexander is a retired pedologist (soils) and active t is timely that Fremontia carry an article concerned with geoecologist who has worked for several agencies, includ- climate change. Systematic observations of biological sys- ing the U.S. Forest Service in California, and investigated I tems are yielding important insights about how they are serpentine soils and plant communities from Baja California responding to climate change. Measuring the ecological effects of climate change requires field observations that to Alaska. span decades, not months or years. So it takes quite awhile before scientists are able to gather sufficient data in order to Jackie Bergquist is the landscape supervisor at Children’s draw meaningful conclusions. Fairyland in Oakland, CA. Jim Bishop’s lead article in this issue is about climate change. In it, Jim introduces us to a fascinating interna- Jim Bishop, with his botanist wife Catie Bishop, has con- tional research effort—the Global Observation Research ducted GLORIA field work since the project began in Cali- Initiative in Alpine Environments (known simply as fornia in 2004. He also helped to develop the strategy and GLORIA). As he explains, alpine environments are particu- methods for the project’s downslope surveys. larly conducive to measuring the biological effects of global climate changes. This is because human disturbance in the Daniel S. Cooper is an independent consultant based in Los alpine zone is generally minimal, alpine environments span Angeles. From 1999 to 2005 he worked for Audubon Cali- nearly all latitudes and elevations, and they include the fornia. The author of Important Bird Areas of California major climate zones of the world. The article then goes on (2004), he is a member of the LA/Santa Monica Mountains to explain more about the alpine zone, and the environ- mental stresses on alpine plants and how they adapt. Chapter of CNPS. The GLORIA project established its first field sites in Europe in 2001, and three years later the first field sites in John Danielsen is a CNPS Fellow. the Western Hemisphere were set up in California. Jim is Stephen W. Edwards is the third director of the Regional part of the California team of scientists and volunteers that is taking surveys at monitoring sites using the standardized Parks Botanic Garden in Berkeley, CA, and has served in that international protocol so the results can be shared globally. position since 1983. While data gathered so far is preliminary, it suggests that plant diversity is increasing in the alpine zone, which Phyllis M. Faber is a past editor of Fremontia (1984–1999), would be expected if average annual temperatures world- and for many years also served as CNPS publications chair. wide are warming. With GLORIA and similar projects, we stand not only to learn about the response of alpine plants Leah Price Hawks was the editor for Ralph Ingols’ books, to climate change, but to greatly expand our knowledge of and is coauthor of Skyline Wilderness Park, Nature’s Gift to high-elevation vegetation in California and worldwide. Napa Valley. —Bob Hass

FREMONTIA (continued on inside backVOLUME cover) 38:4/39:1, OCTOBER 2010/JANUARY 2011