vol. 38, no. 4 and vol. 39, no. 1 • fremontia...volume 38:4/39:1, october 2010/january 2011...

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JOURNAL OF THE CALIFORNIA NATIVE PLANT SOCIETY

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VOL. 38, NO. 4 AND VOL. 39, NO. 1 • OCTOBER 2010 AND JANUARY 2011

FREMONTIA

USING ALPINE FLORA TO MEASURE ECOLOGICALEFFECTS OF CLIMATE CHANGE,SERPENTINE SOILS AND PLANTS,AND OTHER ARTICLES

USING ALPINE FLORA TO MEASURE ECOLOGICALEFFECTS OF CLIMATE CHANGE,SERPENTINE SOILS AND PLANTS,AND OTHER ARTICLES

F R E M O N T I A V O L U M E 3 8 : 4 / 3 9 : 1 , O C T O B E R 2 0 1 0 / J A N U A R Y 2 0 1 1

STAFF (SACRAMENTO)Acting Exec. Director . . . Sue BrittingFinance & Administration Manager .

Cari PorterMembership & Development Coor-

dinator . . . . . . Stacey FlowerdewConservation Program Director . . . . .

Greg SubaRare Plant Botanist . . . . Aaron SimsVegetation Program Director . . . Julie

EvensVegetation Ecologists . Jennifer Buck,

Kendra SikesEducation Program Director . . . Josie

CrawfordAdministrative Assistant . . . . . Marcy

Millett

STAFF (AT LARGE)Fremontia Editor . . . . . . . Bob HassCNPS Bulletin Editor . . . . . Bob HassLegislative Consultant .Vern GoehringEast Bay Conservation Analyst . . . . .

Mack CastermanWebsite Coordinator . . Mark Naftzger

PROGRAM ADVISORSRare Plant Program Senior Advisor . . .

Jim AndreVegetation Program Senior Advisor . .

Todd Keeler-WolfHorticulture Committee Chair . . . . . .

Laura CampCNPS Press Co-Directors . . . . . Holly

Forbes, Dore BrownPoster Program . . . Bertha McKinley,

Wilma Follette

BOARD OF DIRECTORSBrett Hall (President); Lauren Brown(Vice President); Carol Witham (Trea-surer); Laura Camp (Secretary); AtLarge: Ellen Dean, Arvind Kumar, BrianLeNeve, Nancy Morin, Vince Scheidt,Alison Shilling; Chapter CouncilRepresentatives: Orchid Black, SteveHartman

CHAPTER COUNCILDavid Magney (Chair); Larry Levine(Vice Chair); Marty Foltyn (Secretary)

Alta Peak (Tulare) . . . . Joan StewartBristlecone (Inyo-Mono) . . . . . . . . .

Steve McLaughlinChannel Islands . . . . David MagneyDorothy King Young (Mendocino/

Sonoma Coast) . . . . . Nancy MorinEast Bay . . . . . . . . . . . . Bill J. HuntEl Dorado . . . . . . . . . Susan BrittingKern County . . . . . Dorie GiragosianLos Angeles/Santa Monica Mtns . . . .

Betsey LandisMarin County . . Carolyn LongstrethMilo Baker (Sonoma County) . . . . .

Liz ParsonsMojave Desert . . . . . . Tim ThomasMonterey Bay . . . . Rosemary FosterMount Lassen . . . . . . . Catie BishopNapa Valley . . . . . . Gerald TombocNorth Coast . . . . . . . Larry LevineNorth San Joaquin . . . . Alan MillerOrange County . . . . . Nancy HeulerRedbud (Grass Valley/Auburn) . . . .

Joan A. JerneganRiverside/San Bernardino counties . .

Katie BarrowsSacramento Valley . . . Glen HolsteinSan Diego . . . . . . . . David VarnerSan Gabriel Mtns . . . . Orchid BlackSan Luis Obispo . . . . Kristie HayduSanhedrin (Ukiah) . . . . . . . . . Geri

Hulse-StephensSanta Clara Valley . . . Judy FenertySanta Cruz County . Deanna GiulianoSequoia (Fresno) . . . . Paul MitchellShasta . . . . . . . . . . Ken S. KilbornSierra Foothills (Tuolumne, Cala- veras, Mariposa) . . Robert W. BrownSouth Coast (Palos Verdes) . . . . . .

David BermanTahoe . . . . . . . . . . Michael HoganWillis L. Jepson (Solano) . . . . . . . .

Mary Frances Kelly PohYerba Buena (San Francisco) . . . . .

Ellen Edelson

Staff and board listings are as of April 2012.Printed by Premier Graphics: www.premiergraphics.biz

The California Native Plant Society(CNPS) is a statewide nonprofit organi-zation dedicated to increasing theunderstanding and appreciation ofCalifornia’s native plants, and to pre-serving them and their natural habitatsfor future generations.

CNPS carries out its mission throughscience, conservation advocacy, educa-tion, and horticulture at the local, state,and federal levels. It monitors rare andendangered plants and habitats; acts tosave endangered areas through public-ity, persuasion, and on occasion, legalaction; provides expert testimony togovernment bodies; supports the estab-lishment of native plant preserves; spon-sors workdays to remove invasive plants;and offers a range of educational activi-ties including speaker programs, fieldtrips, native plant sales, horticulturalworkshops, and demonstration gardens.

Since its founding in 1965, the tradi-tional strength of CNPS has been itsdedicated volunteers. CNPS activitiesare organized at the local chapter levelwhere members’ varied interests influ-ence what is done. Volunteers from the33 CNPS chapters annually contributein excess of 97,000 hours (equivalentto 46.5 full-time employees).

CNPS membership is open to all.Members receive the quarterly journal,Fremontia, the quarterly statewide Bul-letin, and newsletters from their localCNPS chapter.

VOL. 38, NO. 4, OCTOBER 2010 ANDVOL 39, NO. 1, JANUARY 2011

F R E M O N T I A

Copyright © 2011California Native Plant Society

Disclaimer:

The views expressed by authors publishedin this journal do not necessarily reflectestablished policy or procedure of CNPS,and their publication in this journal shouldnot be interpreted as an organizationalendorsement—in part or in whole—of theirideas, statements, or opinions.

CALIFORNIA NATIVEPLANT SOCIETY

Dedicated to the Preservation ofthe California Native Flora

Bob Hass, Editor

Beth Hansen-Winter, Designer

Brad Jenkins, Cynthia Roye,and Jake Sigg, Proofreaders

CALIFORNIA NATIVE PLANT SOCIETY

MEMBERSHIPMembership form located on inside back cover;

dues include subscriptions to Fremontia and the CNPS Bulletin

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ISSUE DATE : O C T O B E R 2010 A N D J A N U A R Y 2 0 1 1 . PUBL ICAT ION DATE : M AY 2012

MATERIALS FOR PUBLICATIONCNPS members and others are welcome to contribute materials for publicationin Fremontia. See the inside back cover for submission instructions.

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CONTENTSUSING ALPINE FLORA TO MEASURE ECOLOGICAL EFFECTS OF CLIMATE CHANGEby Jim Bishop .................................................................................................................................................. 2CNPS members have helped to establish a network of high elevation monitoring studies to evaluate changein the alpine flora of California that results from climate change.

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

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

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

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

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

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

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

DEVELOPING A SUSTAINABLE MIX FOR SEED GERMINATION USING LOCAL MATERIALSby Jackie Bergquist, Michelle Laskowski, Brianna Schaefer, and Betty Young ........................................... 50A report on research by the Golden Gate National Parks Conservancy to develop a seed germination mix aseffective 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, treelessbiome being studied carefully for evidence of climate change. Photograph by Stu Weiss.

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ur California team,which is working to es-tablish monitoring sitesin the higher mountains

of the state, is part of an interna-tional effort to detect and measurechanges in the alpine flora thatwould be expected to occur as theclimate changes. As an ecosystem ata climate extreme—one that is verytemperature dependent—the alpine

zone is a sensitive indicator of suchchange.

NEED FOR A UNIVERSALALPINE-MONITORINGPROTOCOL

During the past century, scien-tists have observed changes in al-pine flora and in tree line elevation.In Scandinavia, for example, tree

lines have risen by over 500 feet.Where conditions become less harsh,a greater number of species can sur-vive. Species richness of many al-pine peaks has increased, and onone summit in the European Alpsincreased from one to ten speciesover 100 years. That would be ex-pected if the cold alpine elevationsare warming. But until the last tenyears, there had not been a stan-

O

USING ALPINE FLORA TO MEASURE ECOLOGICALEFFECTS OF CLIMATE CHANGE

by Jim Bishop

he early morning is cool and clear, as long shadows of the last and highest-elevation trees stretch

across 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 the plant list.

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.

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dard, universal, and replicable pro-cess for assessing such changes.

In response to the need for bet-ter scientific data and more validcomparisons, the Global Observa-tion Research Initiative in AlpineEnvironments (GLORIA, http://www.gloria.ac.at/) was conceived at theUniversity of Vienna and establishedat field sites in Europe in 2001.GLORIA defines a protocol that canbe applied to alpine summits any-where in the world. The alpine zoneis ideal for detecting biological ef-fects of global climate changes. Hu-man disturbance there is often mini-mal, it spans nearly all latitudes andelevations, and it samples the majorclimate zones of the world—mari-time and continental, tropical, mid-latitude, and polar.

UNDERSTANDING THEALPINE ZONE

The story begins with the ques-tion of why there is an alpine zone.Understanding the environmentalstresses on alpine plants and theiradaptations is the foundation forviewing their responses. The alpinezone is defined as that above theupper tree line—a remarkably con-sistent elevation in a given region,with the trees dropping out com-pletely over elevation increases ofjust feet. Without the absence oftrees there would be no alpine zone.Why should trees meet such a limitto their growth? Answering thisquestion requires understandinghow temperature limits plant cellgrowth, and how it limits treegrowth.

Plant cell metabolism is verytemperature sensitive, slowing to al-most no cell growth at about 42o to46o F. If the daily mean air tempera-tures (the average for the day) aver-aged over the growing season fallsmuch below about 44o F, trees canno longer subsist. That is a globalobservation, consistent over differ-ent taxonomic families, elevations,

growing season lengths, latitudes,and precipitation regimes.

Tree foliage is effectively cooledby the air flow. Tree root zones areshaded by the crowns and remainclose to the average air temperature.So in effect a tree experiences a coldtemperature limit dictated by aver-age air temperature. Average tem-perature in the lower atmosphere

declines steadily with increasingaltitude. Therefore, above tree lineelevation, the growing season is sim-ply too cool to sustain tree growth.

But other types of plants persistin the alpine zone, in spite of coolerair temperatures there—plants withthe same temperature-limited basicmetabolism as trees. How do plantsthat live above the tree line do it?The secret lies in their form andstature, allowing them to be warmeroverall than air temperature wouldindicate.

The low stature and compactform of alpine plants reduce coolingfrom the air flow and allow absorp-tion of sunlight to warm above-ground plant parts. Foliage of alpineplants can be sun-warmed above airtemperature by some 36o F (so thetemperature of an alpine plant mightbe well over 80o F on a 50o F day).The typically sparse distribution ofalpine plants leaves much ground(usually over 50%, often 80% or90%) exposed to solar heating, andtheir forms transmit more heat to

There are currently five areas in Californiawhere alpine plant survey work is beingconducted as part of the internationalGLORIA Project.

A team hiking to the Dunderberg target region in the central Sierra Nevada. Photographby Catie Bishop. All other photographs by the author, except as noted.


the ground than do raised crowns.Consequently their root systemtemperature is above the average ofthat for shaded sites. In effect, eventhough alpine plants grow at a higherelevation than trees, they experiencea warmer microclimate.

As beautifully adapted as theyare, alpine plants still live at theedge of adequate warmth and theybenefit from the lack of competi-tion from the cold-limited trees, sothey are quite vulnerable to tem-perature changes. Alpine plant eco-systems are also dependent on snowdistribution. Snow shelters manyof them from extremes of wintercold and desiccation. Late-seasonsnowfields retard growth until theymelt, and upon melting release aflush of nutrients and water. Alpinevegetation patterns are strongly con-

trolled by the distribution of snow-fields.

It is expected that at least twovariables related to climate changecould significantly affect alpine eco-systems—warmer temperatures, andamounts and patterns of precipita-tion and snowmelt. Warmer tem-peratures can raise the tree line, andwill raise the elevations at which avariety of alpine and subalpine plantscan be more competitive. Less snowor more snow, and changes in pat-terns of snow deposition and meltschedule can also alter alpine plantdistributions.

GLORIA IN CALIFORNIA

GLORIA began in California in2004 due to the tireless efforts ofConnie Millar of the Pacific South-

west Research Station, USFS, andwas the location of the first projectsites in the Western Hemisphere.Millar remains the force behind allof the California projects and is alsoa contributor to other North Ameri-can sites.

Each GLORIA project is orga-nized around a “site” called a targetregion (TR), and includes a groupof three or four summits, spaced inelevation from the highest peaksdown to near tree line. CaliforniaGLORIA summits now number 18,in 5 TRs, from Langley Peak in thesouthern Sierras to the Carson Rangenear Lake Tahoe, and at several lo-cations in the central White Moun-tains. A team of volunteers, students,and agency employees (who havebeen granted release time from theirnormal duties to help) are conduct-

Alpine flowers. CLOCKWISE FROM UPPER RIGHT: Cushion Townsend daisy (Townsendia condensata), Sierra penstemon (Penstemon heterodoxus),Pacific hulsea (Hulsea algida), cushion buckwheat (Eriogonum ovalifolium), White Mountain buckwheat (Eriogonum gracilipes).


Upper edge of bristlecone-limber pine woodland at Patriarch Grove in the White Mountains,1947 and 2005. Dashed line on 1947 photograph indicates road trace corresponding to roadin 2005 photograph. Note the many young bristlecone pines within and beyond the woodlandin the recent photograph. Most of the global warming in this period took placed after 1970.Photographs courtesy of Daniel Pritchett of the White Mountain Research Station.

ing studies of the alpine ecosystemand its changes. CNPS volunteershave been consistent and importantcontributors to the project.

Surveys will be repeated at five-year intervals to measure change inthe alpine flora. The information thatis gathered will be shared globally,as it is at sites in other countriesaround the world.

THE GLORIA SURVEYPROTOCOL

The summit-centered GLORIAprotocol outlines a set of survey plots(successively smaller plots withinlarger ones) that occupy the slopesof the four main compass directions(N, E, S, and W) on each peak. Theplots lie in two elevation bands: theuppermost band takes in the fivemeters (16.4 feet) in elevation justbelow the peak, and the lower bandlies between five and ten meters (33feet) below the peak.

The materials required to con-duct GLORIA surveys have purposelybeen kept low-tech and therefore areusable worldwide regardless of localresources. A compass, tape-measure,and a string-level (a builder’s levelthat is attached to a string) are allthat are needed for surveying. Col-ored string marks the plots, and pinflags have various uses. A one-meter-square quadrat frame with 100 cellsis used for the finest-scale observa-tions. All reference points are de-fined by compass direction and dis-tance from the “high summit point.”

Information collected includeseach species present (includingvoucher specimens on the first sur-vey); its degree of cover (as a mea-sure of each species’ abundance andits influence on available space, light,air, and water); and the cover ofother elements such as rock or soil.The amount of rock and soil indi-cates the potential for expansion ofplant cover, and plant cover maychange as climate changes (see pro-tocol details, page 6).

Enhancements to the basicGLORIA protocol have been devel-oped in California. One of them, the“California method,” is now an op-tion in the international protocol.The observation provides much bet-ter quantitative data on plant coverover an area that samples much ofthe variation on the summit. It cov-ers a larger area than the one meterby one meter quadrats, but is notsubject to the errors of visual esti-mates of plant cover on the largesurvey plots.

The White Mountain ResearchStation is now a GLORIA WorldMaster Site, where many studies ofalpine ecology complement the ba-sic GLORIA surveys. These addi-tional studies encompass extendedplant surveys, tree line and shrubline mapping, small-scale tempera-ture modeling, insect monitoring,alpine meadow flora, and periglacialprocesses (freeze-thaw cycles). A setof weather stations now extendsfrom the base of the White Moun-tains to the highest summit, provid-ing a direct measure of the climatethere over a range of elevations—important information to relate toany observed ecosystem changes.

LOOKING BELOW THESUMMITS

A major addition to the summitsurveys are the “downslope sur-

veys,” observations that cover thealpine zone below the GLORIA sum-mits and provide a more compre-hensive view of the alpine ecosys-tem throughout its entire elevationspan. To date, such surveys havebeen conducted on four majorslopes, collectively spanning the el-evation from White Mountain Peakat over 14,000 feet down into thebristlecone-limber pine woodland at10,800 feet (see protocol details,page 8).

The data gathered from theslopes reveals flora that could po-tentially be moving up toward thesummits in response to warming. Itshows the distribution of each spe-cies by elevation at a given time.Comparing this with the same spe-cies’ elevation distribution at a latertime (for example, ten years later)can produce an extremely usefulmeasure of just how much the ele-vation distribution of that specieshas shifted over time, very possiblyrising as the world warms.

EARLY RESULTS

What have we learned so far?The initial baseline surveys cannotof themselves reveal change. But theyhave provided useful and detailedinformation on the species that arepresent, their relative abundance,and their detailed distributions onsummits and slopes of the alpinezone in California.


ELEMENTS OF THE GLORIA PROTOCOL

he main objective of the GLORIA protocol is to record the plants that grow on the summit, to note roughlywhere 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 outlinedwith colored string, and field workers examineeach plot carefully for the plants that occurthere.

A set of survey plots is laid out on theslopes of the four main compass directions or“aspects” (N, E, S, and W). They lie in twoelevation bands: the uppermost band spans thefive meters in elevation just below the peak,and the lower band lies between five and tenmeters below the peak.

In each summit area section (SAS)—thearea on one aspect and within one elevationband—the plant cover is visually estimated. Inthe 1m x 1m quadrats (outlined with a framecontaining 100 cells), cover is estimated bynoting the occurrence of each species in eachof the cells. Each group of four quadrats iscentered on a main compass direction at the5-meter perimeter.

Over 60 photographs are taken on eachsummit, 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 surveysystem reference points for placing resurveys.

To learn how the actual temperatures and snow cover are changing, a temperature recorder is buried tencentimeters deep on each aspect. Temperature is recorded approximately hourly for several years. The recorderis then dug up, and the records are recovered. The soil-temperature curve has a narrower daily temperaturerange than the surface-temperature curve, but has a similar average. Changes in average air temperature will bereflected in soil temperature. But when snow covers the ground, the soil temperature remains essentially constantat 32o F, and in that way reveals precisely the periods of snow cover.

A diagram of the survey plots as they would appear on the south slope ofthis summit (HSP is the high summit point). Orange marks the perimetersat the 5-meter and 10-meter levels; pink marks the boundaries of thesouth-slope plots; yellow is the California Method diamond; green is thegrid that locates the one-meter quadrats.

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

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AN ADDITIONAL PROTOCOL:THE CALIFORNIA METHOD

10 X 10-meter “diamond” is cen-tered in each aspect on the 5-meter

perimeter. Within the plot, 400 samplepoints set on a half-meter grid produceuseful quantitative data on plant coverageat the mid-scale (surveys prior to 2009used 100 points). Estimating plant covervisually is difficult, especially for plantsthat have low cover values, and such esti-mates are of very limited accuracy. Thequadrat measurements are much moreaccurate than visual estimates, but theycover only small areas. The 10m X 10mareas yield good accuracy over intermedi-ate areas.

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

A


In California, four offive target regions havebeen resurveyed, andtwo of the downslopesurveys in the WhiteMountains have been re-peated. Many of the re-peat surveys show in-creased species richness.The recent downsloperesurveys also showedan increase in the num-ber of annual species(perennial habit is bestsuited to the demandsof the alpine environ-ment, and alpine an-nuals are uncommon).

However, that may be due, in part,to a wetter-than-normal year.

Findings over the ten years from1994–2004 from the European Alpshave shown reductions in cover ofthe highest elevation species (thosein the zone of permanent snow, orthe “nival” zone), with lesser reduc-tions and some increases within thealpine subzones below that. Thesechanges show that the highest plantsare losing ground to plants fromlower elevations, at least over a re-cent decade.

Even if the changes observed inthe GLORIA resurveys are signifi-cant, it is not possible to attributechange over five years to long-termclimate trends. Inter-seasonal varia-tion can easily affect the flora rep-resented in a given year. The valueof the current comparisons is thatthey demonstrate the potential of

DOWNSLOPE SURVEY PROTOCOL

ownslope surveys are conducted on some of the slopes belowthe GLORIA summits, into the subalpine woodland. Meter-

wide belt transects along the elevation contour are spaced every 25meters (82 feet) in elevation. Each transect is 100 meters long (100square meters total area), divided into 10-meter segments. All spe-

cies present in each 10-meter segment are recorded. Atsample points spaced every half meter (400 for theentire transect) the plant species or other cover type isidentified. The downslope transects duplicate, in areaand sampling density, the 10m X 10m California-method plots on the summits.

Outline of one segment, 1 meter wide and 10 meters long, centered on the whitetape, of the 100-meter belt transect at that elevation. All species are noted withinthe segment. Sample point positions for plant species or cover type are indicatedon this photograph with red circles. “Frequency” refers to the total number ofsegments a species occurs in; “Cover” refers to the number of sample points thatcontact a given species or cover type.

LEFT: A pointer that marks two sample points, one at each end, ismoved along and placed every one-half meter along the tape. •BELOW: Botanists look closely for small plants, since it is easy tomiss some of the Alpine species, which can be quite tiny.

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the protocols to reveal change.However, only changes in a direc-tion consistent with a warmingworld, sustained over two or moredecades, will indeed be compellingevidence.

Current data is also being com-pared to earlier botanical data fromthe White Mountains. For example,the uppermost occurrence on gra-nitic substrates of dwarf sagebrush(Artemisia arbuscula) as reported in1961 has been compared to the re-cent upper shrub line surveys anddownslope surveys. Based on thoseobservations, which span a 50-yearinterval, this species’ upper limit has

T he team is finishing up its work on the summit. Strings are wrapped on cardboard holders, data

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.

risen 500 feet, and several commonalpine plants have been reduced incover wherever the Artemisia hasencroached. Aerial photo compari-sons from the late 1940s into the2000s show bristlecone pine wood-land filling in along its upper mar-gins and young trees spreadingupslope.

Much remains to be learned, andonly time will reveal long-termchange. But the baseline data beingestablished, and the early look atchanges, are an important contribu-tion to assessing the impacts of cli-mate change on the alpine ecosys-tem across the world.

REFERENCES

Koerner, C. 2003. Alpine Plant Life,Functional Ecology of High MountainEcosystems, 2d. ed. Springer-Verlag,Berlin, Heidelberg, New York.

IPCC Working Group 1. 2007. ClimateChange 2007, the Physical ScienceBasis. Cambridge University Press,Cambridge, MA.

White Mountain Research Station,GLORIA website: http://www.wmrs.edu/projects/gloria%20project/default.htm

Jim Bishop, 1144 Mount Ida Road,Oroville, CA 95966, [email protected]

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MANAGING A MOUNTAIN: THE SAN BRUNOMOUNTAIN HABITAT CONSERVATION PLAN

by Patrick Kobernus

conserved habitat.1 As of 2012, mostof the development on the moun-tain has been completed.

Management of invasive speciesto protect the endangered specieshabitat on the mountain has beenlargely successful over the past 30years. However, coastal scrub suc-cession, in combination with ex-

panding populations of invasive spe-cies, continue to overtake grasslandhabitat on the mountain. The futureconservation of the endangered spe-cies and their habitats, now morethan ever, depends upon implemen-tation of a more comprehensive habi-tat management program to protectthese species for future generations.

MOUNTAINFLORA ANDFAUNA

San Bruno Mountainis located in northernSan Mateo County, ad-jacent to San Francisco.It consists of approxi-mately 2,830 acres ofopen space, and is bor-dered by the urban andsuburban portions ofDaly City, South SanFrancisco, Colma andBrisbane. Though it isisolated by urbanization,the mountain is consid-ered the northernmostpart of the Santa CruzMountains.

The famed botanistJames Roof asserted thatSan Bruno Mountainsupports one of the lastand the most expansiveareas of a unique andhighly diverse grasslandand shrubland flora,which he referred to as“Franciscan” (Edwards2000). This Franciscan

n 1982, the first Habitat Conserva-tion Plan (HCP) in the nation wasapproved for San Bruno Moun-tain. Over the past 30 years most

of the mountain, approximately2,830 acres (82%), has been con-served as habitat for three endan-gered butterfly species, wildlife, andplants; and approximately 350 acres(10%) of the mountain has been de-veloped. An additional 260 acres(8%) of land remains undeveloped,and it is likely that a majority of thisland will ultimately be set aside as

The San Bruno Mountain Habitat Conservation Plan—adopted in 1982 and the first HCP in the nation—is responsible for protecting host plants for three endangered butterflies, and conserving the area’sdiverse native flora and fauna. All photographs by the author unless otherwise indicated.

I1 Most of the remaining “unplanned parcels” will likely be conserved due to these lands

being located on very steep slopes with no infrastructure (roads, utilities) for development.

The HCP requires that a minimum of 40% of this land be dedicated as conserved habitat.

F R E M O N T I A 1 1V O L U M E 3 8 : 4 / 3 9 : 1 , O C T O B E R 2 0 1 0 / J A N U A R Y 2 0 1 1

flora was once common through-out the hills of San Francisco buthas been almost entirely destroyedin the city by development andplanting of nonnative trees.

The grassland on San BrunoMountain is actually a combina-tion of different types of grass-lands intergrading and sharingsome of the same wildflower andshrub associates. Grassland typesvary on the mountain de-pending on elevation, ex-posure, and soil type. Thedryer southern exposurestend to have stands ofpurple needle grass (Nas-sella pulchra), while themore fog-shrouded grass-lands near the summitare dominated by Califor-nia fescue (Festuca califor-nica), red fescue (F. rubra),and Idaho fescue (F. ida-hoensis). There are alsostands of California oatgrass (Danthonia califor-nica), blue wild rye (Ely-mus glaucus var. glaucus),Pacific reed grass (Calamagrostisnutkaensis), June grass (Koeleriamacrantha), and tufted hair grass(Deschampsia cespitosa ssp. holci-formis).

As a biologist hired to moni-tor the three endangered butter-fly species on San Bruno Moun-tain for 13 years, I can attest to itsunique beauty. Each spring andsummer, I would hike the moun-tain repeatedly while I recordedmy observations of the mission blue,San Bruno elfin, and callippesilverspot butterflies, and the statusof their grassland habitats.

The San Bruno elfin’s host plant,Pacific stonecrop (Sedum spathuli-folium), grows in coastal prairie andon rocky outcrops and roadcuts.The Callippe silverspot’s host plant,Johnny jump up (Viola pedunculata),grows in coastal prairie and in non-native annual grasslands. The mis-sion blue’s host plants, silver lupine(Lupinus albifrons var. collinus), var-

ied-colored lupine (L. variicolor),and summer lupine (L. formosus var.formosus) grow within coastal prai-rie, nonnative annual grassland,

rocky outcrops, roadcuts, and oncut slopes.

The mountain is not onlyhome to three endangered butter-fly species but also supports a widediversity of other native flora andfauna. Many varieties of wildflow-ers can be found on the moun-tain, including coast rock cress(Arabis blepharophylla), Pacificstonecrop, varied-color lupine,

Johnny jump up, gold-fields (Lasthenia cali-fornica), shooting stars(Dodecatheon hendersonii),blue larkspur (Delphin-ium decorum), farewell tospring (Clarkia rubicun-da), and owl’s clover (Cas-tilleja densiflora), amongmany others. Each patchof grassland is a uniquelybeautiful “natural garden”that has been constructedthrough the forces of na-ture and time. Each Marchthrough June, the grass-lands and wildflowersemerge and change into

new combinations of color andbeauty as the season progresses.

There are several rare plantspecies including two that arestate and/or federally listed, SanBruno Mountain manzanita (Arc-tostaphylos imbricata) and SanFrancisco lessingia (Lessingiagermanorum). There are also sev-eral California Rare Plant Rank1B species (formerly CNPS List1B) such as Montara manzanita

(Arctostaphylos montaraensis), Paci-fic manzanita (Arctostaphylos pa-cifica), Diablo helianthella (Helian-thella castanea), San Francisco spine-flower (Chorizanthe cuspidata var.cuspidata), and San Francisco cam-pion (Silene verecunda ssp. verecun-da). Other rarities include an arach-nid, incredible harvestman (Bank-sula incredula); a solitary bee (Du-fourea stagei); and several range-limited endemic plants.

Plant communities on the moun-tain include northern coastal scrub,

TOP: A freshly emerged male mission bluebutterfly (Icaricia icarioides missionensis),found in San Bruno Mountain’s grasslandhabitat. It is an endangered species. •MIDDLE: A mission blue larva feeding onsilver lupine (Lupinus albifrons var. collinus)on the west peak of San Bruno Mountain.• BOTTOM: A San Bruno elfin larva foragingon the flowerheads of its host plant, Pacificstonecrop (Sedum spathulifolium) near thesummit of San Bruno Mountain.


coast live oak woodland, coastal ter-race prairie, freshwater marshes andseeps, central coast riparian scrub,nonnative gorse and broom scru-blands, nonnative eucalyptus for-est, and nonnative annual grass-land. The most dominant vegeta-tion on the mountain is northerncoastal scrub and nonnative annualgrassland.

FIRST HABITATCONSERVATION PLANIN U.S.

Since 1982 the mountain hasbeen the site of the first HabitatConservation Plan (HCP) in thenation. HCPs were created as amechanism to balance private prop-erty rights and endangered speciesprotection, by allowing limited “tak-

ing” (destruction) of endangeredspecies and their habitat, providedthat the taking will not appreciablyreduce the likelihood of the sur-vival and recovery of the species inthe wild, as specified in section10(a)(1)(B) of the federal Endan-gered Species Act.

This mechanism has been usedas a tool for settling land disputesby allowing landowners to econo-mically develop portions of theirproperties, while simultaneously en-suring long-term protection of en-dangered species through dedicationof conservation areas, habitat man-agement and monitoring, and/orother mechanisms. As of 2010, over700 HCPs have been permitted inthe U.S. by the U.S. Fish and Wild-life Service (USFWS 2010).

The primary purpose of the SanBruno Mountain HCP is to protectthe grassland habitat that supportsthe three endangered butterfly spe-cies, while allowing limited devel-opment to occur. Prior to the for-mation of the HCP, approximately1,950 acres of land on San BrunoMountain had been purchased and/or donated to create the San Bruno

Mountain State andCounty Park. Thisland contained vir-tually the entirehabitat for the SanBruno elfin butter-fly on the mountain.However, it did notinclude the primehabitat areas for themission blue andcallippe silverspotbutterflies, whichwere located on theeastern portions ofthe mountain.

The HCP pro-vided a mechanismby which an addi-tional 800+ acresof habitat wouldbe conserved andadded to the Park,and all of the con-

served land within the park wouldbe managed and monitored for theendangered species as well as forother native flora and fauna. Withinthe current 2,830-acre conservationarea, approximately 90% of the mis-sion blue and callippe silverspot but-terflies’ habitat, and 100% of the San

Bruno elfin butterfly’s habitat hasbeen protected.

The development permittedthrough the HCP is primarily rele-gated to the lower slopes of themountain, thereby protecting themajority of higher quality butterflyhabitat on the upper ridges and hill-tops. For the callippe silverspot, thiswas critical, because this species re-quires hilltops for mate selection.Male callippes stake out territorieson the highest hilltops or ridgelinesavailable to encounter and attractfemales for mating.

The conservation areas also in-cluded protection of several rareplant species, with the exception ofmost of the San Francisco lessingiaand San Francisco spineflower pop-ulations, which are located on pri-vate property on the west side ofthe mountain. These populations arestill extant, however developmentand invasive species are potentialthreats.

The HCP specified the impor-tance of management and monitor-ing of the butterflies’ habitat andprovided a funding mechanism tosupport these activities—an annualmonetary assessment on every resi-dence and business that is builtwithin the HCP boundary. This as-sessment rises with the annual in-flation rate, and has provided a con-sistent level of funding since the cre-ation of the HCP.

The HCP fund is managed bythe HCP Trustees (the City Manag-ers of Daly City, Brisbane, and SouthSan Francisco, and the San MateoCounty Manager). Monitoring andhabitat management is implementedby San Mateo County Parks Depart-ment, and implementation of theplan is overseen by the U.S. Fishand Wildlife Service.

THREATS FROM INVASIVESPECIES

In 1982, an assortment of ag-gressive invasive plant species were

View of housing development and mission blue butterfly hostplant, silver lupine (Lupinus albifrons var. collinus) on thesoutheast ridge of San Bruno Mountain. Development is nolonger the most serious threat to the grassland habitat on themountain. A greater threat is coastal scrub succession and theexpansion of invasive species.


identified and mapped on the moun-tain. Gorse (Ulex europaeus), themost aggressive of these plants, wasintroduced to the mountain in the1930s and by 1982 had expanded tocover several hundred acres of themountain. Based on the rate of ex-pansion of this plant and that ofother invasive weeds, combined withillegal off-road vehicle use andcoastal scrub succession, it was esti-mated that the habitat for the en-dangered butterfly species on themountain could be completely wipedout within 50 to 200 years (SanBruno Mountain HCP Steering Com-mittee 1982).

As a result, the funding and

implementation of ongoing habitatmanagement with an emphasis oncontrolling invasive species becamean important component of the SanBruno Mountain HCP. Due to thelack of information on the feasibil-ity of habitat restoration at the timeof the inception of the HCP, theHCP’s primary goal has been focusedon the conservation and manage-ment of existing habitat for the but-terflies (San Bruno Mountain HCPSteering Committee 1982).

Though the HCP has often beencriticized for the lack of restorationwork that has been conducted, thestrategy of focusing efforts on pro-tecting the existing habitat has

proven to be successful in maintain-ing most of the habitat for the en-dangered species over the 30-yearspan of the HCP. While the reduc-tions of large infestations has beeneffective, it is the less noticeable habi-tat maintenance work that is doneyear in and year out by HCP workcrews and volunteer groups thatserves to protect the majority of thebutterfly habitat on the mountain.

HABITAT RESTORATION

Restoration of mission blue andcallippe silverspot habitat is re-quired by the HCP within areas thatwere disturbed by grading activi-

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


ties adjacent to the developments.These areas have steep slopes thatwere engineered to protect againstlandslides. Restoration of butterflyhabitat has not been successful onmost of these slopes, due to thedifficulty in establishing nativeplants in poor soil conditions. Thisis especially true for the host andnectar plants for the callippesilverspot butterfly. Propagating andreplanting of the callippes’ hostplant Johnny jump up has shownto be expensive, with very little suc-cess to date. Johnny jump up isdifficult to grow under nursery con-ditions, and has had a very lowsurvival rate when transplanted intorestoration sites. For the missionblue however, its host plants (espe-cially silver lupine) have been re-established on several restorationslopes because these plants areadapted to disturbed rocky slopeswith thin or poor soil conditions.

CURRENT THREATS ANDMANAGEMENT

While efforts have been success-ful in reducing the large, woody in-vasive species on San Bruno Moun-tain, control work has been less ef-fective at stemming the tide of coastalscrub succession. An independentanalysis of almost 20 years of but-terfly data collected over the courseof the HCP revealed that the overalldistribution of the mission blue andcallippe silverspot butterflies re-mained stable. However, geographicareas of concern were identified foreach species.

For the period between 1982 and2004, San Bruno Mountain lost anestimated 122 acres (8.6%) of grass-land habitat. This was primarily dueto coastal scrub succession withinthe HCP conservation area (San

Mateo County Parks Department2008). This corresponds to a loss ofover five acres of grassland per year.The expansion of coastal scrub veg-etation and corresponding loss ingrassland has been documented inmany regions of California (Murray2003, McBride and Heady 1968),and is often the result of the re-moval of grazing and/or burningfrom a grassland ecosystem.

Historically cattle grazing andbrush burning by local ranchers re-sulted in the control of coastal scrubvegetation on San Bruno Mountain,but also facilitated the spread of in-vasive plant species. Invasive grassessuch as ripgut brome (Bromus dian-drus), velvetgrass (Holcus lanatus),and invasive herbaceous weeds suchas fennel (Foeniculum vulgare), wildradish (Raphanus sativus), and oxa-lis (Oxalis pes-caprae) have prolifer-ated because of the ability of thesespecies to rapidly expand into grass-lands (San Mateo County ParksDepartment 2008). Atmosphericsources of nitrogen pollution (smog)may also be contributing to thespread of these invasive grasses andweeds within the grasslands (Weiss2006).

As more and more weeds prolif-erate and die back, the resultantaccumulation of live and dead bio-mass (thatch) reduces the amountof light reaching the soil surface,suppressing the growth of nativegrassland plants. Increased mois-ture retention from the shade cre-ated by thatch may also facilitatethe expansion of coastal scrub intothe grassland areas. Furthermore,where invasive control work hasbeen done for decades, there is asignificant build-up of thatch fromold stalks of fennel, broom, andgorse plants that were left to decayin place.

The level of thatch within the

grasslands on San Bruno Mountainwas evaluated in 2002 using liveand dead above-ground biomassmeasurements. Values measuredwithin the grasslands on the southslope of the mountain prior toexperimental grazing treatments,showed live and dead above groundbiomass levels of 5,000 to 9,000lbs/acre. A large proportion of thiswas from thatch. As a comparison,the recommended ranges for Re-sidual Dry Matter (live biomass) incoastal prairie grasslands with mini-mal woody plant cover ranges from1,200 to 2,100 lbs/acre (UC Davis2002).

The reduction in wildfires, re-moval of grazing animals in the early1960s from the mountain, and at-mospheric nitrogen pollution are alllikely factors contributing to the pro-liferation of invasive plants, build-up of thatch, and brush successionon the mountain.

FUTURE MANAGEMENT

The San Bruno Mountain HCPhas been an experiment—the firstof its kind—to protect endangeredspecies habitat while allowing lim-ited development. For 30 years theplan has been a qualified success inthat all three of the endangered but-terfly species on the mountain con-tinue to be locally abundant. How-ever, management of the conserva-tion areas will need to adapt tochanging conditions and addressproblems such as coastal scrub suc-cession and invasive weeds in a morecomprehensive way. The 2008 SanBruno Mountain Habitat Manage-ment Plan spells out in detail thepriority areas to protect, the currentand emerging threats to the moun-tain’s habitats, and the methods formonitoring and management to ad-dress these threats.

FACING PAGE, TOP: Buckeye Canyon in 1986. This photograph was taken approximately 25 years after cessation of cattle grazing on SanBruno 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 EnvironmentalSciences and San Mateo County Parks Department.


Until 2010, threats to nativehabitats on San Bruno Mountaincould not be addressed comprehen-sively given the existing manage-ment budget. However, as a resultof an agreement reached betweendevelopers, the city of Brisbane, SanMateo County Parks Department,and the U.S. Fish and Wildlife Ser-vice, an additional four million dol-lars will be generated through de-velopment fees and placed into anendowment for the mountain. Oncecollected, these funds would increasethe HCP annual budget for habitatmanagement by two to three timesits former level. These funds need tobe used to manage more grasslandareas on the mountain.

With accelerated changes ex-

pected to occur from global climatechange, it is important to preserveas much potential grassland habitatas possible to buffer the endangeredspecies from occasional large-scaledeclines in habitat quality. For ex-ample, in the extremely wet El Niñoyear of 1998, the numbers of mis-sion blue butterflies on San BrunoMountain declined markedly in ar-eas where silver lupine was the domi-nant host plant. Silver lupine expe-rienced a widespread die-off due toa fungal infestation brought on bythe excessive wet soil conditions.This impact was observed through-out the range of the species (includ-ing the Marin headlands and at TwinPeaks, San Francisco).

In contrast, habitat areas on San

Bruno Mountain that supported thealternative host plant summer lu-pine were unaffected by the fungus,and mission blue butterfly numbersin these areas remained stable. Inthe subsequent 14 years, silver lu-pine plants have rebounded signifi-cantly, as have the mission bluenumbers in the areas impacted bythe fungus. Protecting areas of dif-ferent habitat quality, slope aspect,and within different microclimatesis important because habitat qualitycan be expected to fluctuate overtime, due to plant senescence andclimatic factors, and these fluctua-tions may become more extreme inthe future.

While prescribed burning maycontinue to be difficult to imple-

View of the ridge above Owl Canyon near the summit of San Bruno Mountain. The area contains prime grassland habitat, including nativewildflowers and shrubs, for the Callippe silverspot and San Bruno elfin butterflies, both endangered.


ment on San Bruno Mountain due topublic safety concerns, grazing and/or mowing should be implementedto reduce vegetation fuel loads be-tween parkland areas and homes andbusinesses. Stewardship grazing ormowing of 100–500-foot bufferzones on regular intervals would re-duce fuel loads near populated areasand could potentially allow for thesafe use of controlled burns in someareas of the mountain. Also, grazingor mowing within lower elevationareas between parklands and devel-opments would not impact the moreintact stands of coastal prairie, whichare more concentrated on the upperslopes of the mountain.

Grazing, mowing, and/or burn-ing will need to be applied to ad-dress scrub succession and invasiveweed infestations on San BrunoMountain. These tools will need tobe used in combination with otherweed control methods to manageareas effectively. There must be aprescriptive approach that is tailoredin timing, duration, and frequencyto each area of the mountain, de-pending upon the grassland type,surrounding terrain, presence of rareand endangered species, and publicsafety concerns.

CONCLUSIONS

Over the years, the San BrunoMountain HCP has received a sub-stantial amount of criticism fromenvironmental groups regarding thelack of successful habitat restora-tion on the mountain. In contrast,resource managers have emphasizedthe positive aspects of the HCP andhow it has worked to protect theendangered species habitat and na-tive plant communities. The realityis that both groups are right. Therestoration work has been largelyunsuccessful on the graded slopes,while the habitat management hasbeen successful in protecting theendangered species populationswithin the conservation areas.

The primary focus of environ-

mental groups has been on fightingdevelopment on San Bruno Moun-tain, under the assumption that de-velopment is the major threat tothe endangered species. In reality,though, the permitted developmenthas impacted approximately 10%of the mountain and was relegatedto lower slopes, generally of lesserhabitat value. The development isnow nearing completion. The onlyway to protect the endangered spe-cies and the plant communities ofSan Bruno Mountain for future gen-erations will be to manage the re-maining conserved habitat moreeffectively.

The 2008 Habitat ManagementPlan for San Bruno Mountain estab-lished a goal of maintaining between1,200–1,800 acres of native and non-native grassland on the mountain.Currently the area of grassland isapproximately 1,250 acres, but it isdecreasing at a rate of approximatelyfive acres per year. Slowing the rateof coastal scrub succession and in-creasing the amount of grassland willrequire that brush control programsbe implemented sooner rather thanlater.

The San Bruno Mountain storyis not unique: brush succession andinvasive species have been negativelyimpacting grasslands and meadowsthroughout California for severaldecades. The San Bruno MountainHCP is unusual, however, in that ithas had a mandate and funding toaddress these issues for 30 years. Itwill take a coordinated effort on thepart of biological monitors, habitatmanagers, oversight agencies, envi-ronmental groups, and the commu-nity to work cooperatively and cre-atively to ensure that the mountain’sendangered species and uniqueFranciscan flora are protected forthe next 30 years.

REFERENCES

Edwards, S.W., 2000. The FranciscanRegion. Transcript of a Lecture by JamesRoof ca. 1975. The Four Seasons,

Journal of the Regional Parks BotanicGarden, Volume 11, Number 2. Oc-tober 2000. Published by East BayRegional Park District, Oakland, CA.

McBride, J., and H.F. Heady, 1968. In-vasion of Grasslands by BaccharisPilularis. Journal of Range Manage-ment. 21:106-108.

Murray, M.P., 2003. Losing ground: wil-derness meadows and tree invasionover a 55-year period in California’sKlamath Range. Pp. 81-97, In: K.L.Mergenthaler, J.E. Williams, andE.S. Jules (eds). Proceedings of theSecond Conference on Klamath-Siskiyou Ecology. Siskiyou FieldInstitute, Cave Junction, OR.

San Bruno Mountain Habitat Conser-vation Plan Steering Committee,1982. San Bruno Mountain HabitatConservation Plan, Volumes I and II.Chaired by the County of San Mateo.Plan prepared by Thomas Reid As-sociates, November.

San Mateo County Parks Department,2008. San Bruno Mountain HabitatManagement Plan, 2007. Preparedin Support of the San Bruno Moun-tain Habitat Conservation Plan. Re-vised 2008. San Mateo County Parksand Recreation Division. March2008.

University California at Davis, 2002.California Guidelines for ResidualDry Matter (RDM) Management onCoastal and Foothill Annual Range-lands. Rangeland Monitoring Series,Publication 8092. University of Cali-fornia Davis, Division of Agricultureand Natural Resources, CaliforniaRangelands Research and Informa-tion Center.

U.S. Fish and Wildlife Service, 2010.U.S. Fish and Wildlife Service, Con-servation Plans and Agreements Da-tabase. Website accessed October 10,2010. http://ecos.fws.gov/conserv_plans/public.jsp

Weiss, S.B., 2006. Impacts of nitrogendeposition on California ecosystemsand biodiversity, California EnergyCommission Report. Website publi-cation accessed on June 25, 2007.http://www.creeksidescience.com/publications.html

Patrick Kobernus, 1072 Geneva Avenue,San Francisco, CA 94112, [email protected]


RARE PLANTS OF GRIFFITH PARK, LOS ANGELESby Daniel S. Cooper

ising more than a thousand feetoff the floor of the Los Ange-les Basin, Griffith Park dividesthe eastern San Fernando Val-

ley from the coastal plain of LosAngeles and protects more than4,000 acres of rugged slopes andcanyons at the eastern end of theSanta Monica Mountains. Althoughportions of the park are heavily used,with golf courses, grassy picnic ar-eas, and other attractions situatedmainly at the park’s borders, its rug-

ged interior is still surprisingly wild,owing to both its steep topographyand lack of established trails.

WILDFIRE PROVIDESIMPETUS FOR NEWRESEARCH

On May 8, 2007, an arson-caused wildfire burned about 20%of the park, essentially the entiresoutheastern corner, threatening

such landmarks as the Griffith Ob-servatory and the Los Angeles Zoo,none of which sustained damage.Just a few weeks earlier, on March30, an arson fire had swept up to-ward the northwestern corner of thepark, burning a large area of high-elevation chaparral atop CahuengaPeak, the highest point in the east-ern Santa Monica Mountains. Whilethese fires radically altered portionsof the park’s vegetation—at leasttemporarily—their more lasting

View southeast from block F8 of Griffith Park (see Figure 2) showing downtown Los Angeles (at right) and the Los Angeles River (center).

R


contribution may have been the in-creased awareness they elicited fromlocal conservationists and city staffalike, most of whom had never ex-plored the park. Suddenly, the parkseemed like a worthy subject of im-mediate ecological exploration, per-haps because it was almost com-pletely reduced to ash-coveredslopes.

Just prior to the fires and con-tinuing to the present, a collectionof neighborhood groups and the LosAngeles/Santa Monica MountainsChapter of CNPS funded severalprojects aimed at gathering baselinebiological data on its plants and ani-mals. These were carried out in con-junction with the development of awildlife management plan fundedby the City of Los Angeles (Cooperand Mathewson 2009). While theflora of the Santa Monica Moun-tains as a whole has been well docu-mented (Wishner 1997; Gibson andPrigge 2003; California Native PlantSociety, undated), that of the fareastern end of the range is compara-tively less well-known than the large,protected areas to the west such asTopanga Canyon State Park and theSanta Monica Mountains NationalRecreation Area.

Aside from scattered references(e.g., McAuley 1985), the flora ofGriffith Park had never been syn-thesized, although dozens of speci-mens had been collected here sincethe mid-1800s, mostly prior to 1950(Consortium of California Herbaria2011). The one available checklistof the plants of the park (Brusha2003) does not differentiate betweenplanted and naturally occurringtaxa, and presents just a fraction ofthe true species diversity. Currentlyan as-yet unpublished checklist ofthe park’s flora stands at more than500 species vouchered or photo-graphed (Cooper, forthcoming).This is nearly half the known floraof the Santa Monica Mountains andincludes 350 natives. Still, the parkhas apparently never been a popu-lar place for botanizing, probably

because of its rugged topographyand the fact that most readily acces-sible hiking areas (mainly fire roads)have been degraded by weed abate-ment practices and non-native veg-etation.

SURVEY LAUNCHED

To remedy this information gapand to identify critical areas for plantconservation, an informal, volun-teer-based “Griffith Park Rare PlantSurvey” was launched in spring 2010(Cooper 2010). The botanical his-tory of an area can be a challenge touncover, even for trained research-ers. Most parts of California remainvery poorly known botanically, in-cluding sites within urban areas. Of-ten there are a few local experts whohave botanized in an areaand have kept at leastsome field notes (or to-day, digital photographs),and can help put togethera list of species likely tooccur based on their ex-perience in the region.Such observations maybe supplemented by plantcollections in local her-baria, which are often theonly record of specieslong gone from urbansites such as native wild-flowers.

Through my ownfieldwork conductingwildlife surveys in thepark since 2007, I wasable to identify key habi-tat features most likelyto hold rare or interest-ing plants, such as rockoutcrops and remotecanyons. To begin thetask of locating rare andsignificant species andpopulations, a list of tar-get species was devel-oped with the help of RichardFisher, Bart O’Brien, and CarlWishner, local botanists familiarwith the flora of the Santa Monica

Mountains. Included in this list wereboth CNPS rare taxa (CaliforniaNative Plant Society 2010), as wellas a group of species we felt weresignificant, either because they arerare in the region, or because iffound, they would represent dis-junct occurrences from populationsfarther west in the Santa MonicaMountains, or to the north in theVerdugo and/or San Gabriel Moun-tains.

I also searched for taxa knownin the area only from historical col-lections (Consortium of CaliforniaHerbaria 2011), though some ofthese were likely collected in areasno longer undeveloped (e.g., the LosAngeles River and “ProvidenceRanch,” since converted to ForestLawn Memorial Park). A small group

of volunteers used this species list,along with a map of the park(Cartifact, Inc. 2007) that had beendivided into 40 survey blocks, and

Figure 1. Locations of Rare Plants inGriffith Park

KEY:Green: Nevin’s barberry (presumed planted)Magenta: Plummer’s mariposa lily (generalized

locations; plant is very widespread)White: Catalina mariposa lilyYellow: Slender mariposa lilyDark Blue: Clay bindweedLight Orange: Humboldt lilyLight Blue: Hubby’s phaceliaRed: San Gabriel Mountains leather oak (pending

further identification)


sent me photos and notes from theirforays throughout spring 2010.

RARE SPECIES FINDINGS

From this effort, 13 rare species(California Rare Plant Ranks 1, 2,and 4) were identified as occurringor having occurred in or adjacent toGriffith Park. Of these, nine werefound to be extant (still in exist-ence), and four are known only from

historical specimens (Table 1). (Adescription of the California RarePlant Ranks, formerly the CNPSLists, is available at http://www.cnps.org/cnps/rareplants/ranking.php.) Of the four species believedextirpated from Griffith Park, onlyone, the Brewer’s redmaids (Calan-drinia breweri), has been collectedrecently in the region near the park(Verdugo Mountains); the remain-ing species are apparently very rare

in the Los Angeles area if they per-sist at all.

Three species of mariposa lilyoccur in the park, and exhibit aninteresting, non-overlapping patternof occurrence, with Plummer’s mari-posa lily (Calochortus plummerae)by far the most common. Hundredsof these spectacular pink and yellowlilies were encountered on thin, grav-elly soil along ridges, often in asso-ciation with yucca (Hesperoyuccawhipplei), giant stipa (Achnatherumcoronatum), coast buckwheat (Erio-gonum fasciculatum), and chamise(Adenostoma fasciculatum). Theywere often found along small trailsand footpaths (including equestrianroutes), where this activity appearedto limit non-native grasses, amongother plants, that might otherwiseovertake the lilies.

Catalina mariposa lily was foundin pockets of grassland on moist,heavy clay soil near the south-cen-tral area of the park, with two largepopulations discovered that exceed100 plants. Finally, the slender mari-posa lily (Calochortus clavatus var.gracilis) was discovered prior to thestart of the survey, in a single patchof fewer than 10 individual plants ina grassy opening in chaparral on aremote peak near the center of thepark. No other populations of thisstunning yellow lily were encoun-tered, despite much searching.

Humboldt lily (Lilium humboldtiivar. ocellatum) appears to be con-fined to just four oak-shaded can-yons in the park, with most plants(50+ individuals) located in Brush

Outline of park is drawn in gray. This grid system follows that used by Cartifact, Inc.(2007) for a Griffith Park map, and notable natural and physical features are listed withineach block. The colors correspond to the number of rare species found within each block,as indicated by the key on the right. For example, pale pink blocks had two to three rarespecies, while the darkest red blocks had nine or more.

Figure 2. Distribution of Rare and Locally Rare Plant Species inGriffith Park, by Survey Block

LEFT: Plummer’s mariposa lily is found frequently on steep ridges on eroding soil throughout Griffith Park, Los Angeles. All photographsby 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.


Species Legal statusRepresentative Park statusspecimen1 (40 survey blocks)

Nevin’s barberry Federal: SBBG37272 Probably introduced;Berberis nevinii Endangered found in 5 survey

State: blocksEndangeredCA Rare PlantRank 1B.1

Brewer’s redmaids CA Rare Plant JEPS17234 HistoricalCalandrinia 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 plantsvar. 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 HistoricalDudleya multicaulis Rank 1B.2 specimen(s); no

modern record

large-leaved filaree CA Rare Plant RSA390952 HistoricalErodium 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 mappedJuglans californica

Humboldt lily CA Rare Plant (photograph 6 survey blocks,Lilium humboldtii Rank 4.2 only) mesic canyonsvar. 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 HistoricalPiperia cooperi Rank 4.2 (“Providencia Ranch”) specimen(s);

no modern record

San Gabriel Mountains CA Rare Plant RSA652868 1 survey blockleather oak Rank 4.2Quercus durata var. gabrielensis

Sources: Consortium of California Herbaria 2011; Cooper 2010.

1 Location is “Griffith Park” unless otherwise indicated.SBBG = Santa Barbara Botanic GardenJEPS = Jepson Herbarium, UC Berkeley

RSA = Rancho Santa Ana Botanic Garden HerbariumLA = UCLA HerbariumUCR = UC Riverside Herbarium

TABLE 1. SPECIAL STATUS PLANTS AT GRIFFITH PARK


Canyon in the southwest corner.Hubby’s phacelia (Phacelia cicutariavar. hubbyi) was found on exposed,eroding sedimentary deposits at thefar southeast corner of the park, onsoil probably deposited by the an-cient Los Angeles River that flowsnearby. Thousands of these plantsbloom in early spring in the under-story of open black walnut wood-land. The tiny clay bindweed (Con-volvulus simulans) was found by avolunteer in 2010 on a moist, grassyslope on heavy clay near the south-western border of the park just northof Hollywood, a site that also sup-ports the only park population ofchocolate lily (Fritillaria biflora).

Nevin’s barberry (Berberis nevinii)is arguably the most famous rare plantin the park, and also the most prob-lematic. Most stands (dozens ofplants each) occur in two popula-tions. Scattered plants located since2007 have all been along major roads(strongly suggesting a horticulturalorigin), or just down-slope of estab-lished populations. Whether plantsat Griffith Park originated from lo-cally-collected seed decades ago—possibly from now-extirpated nativepopulations in the Los Angeles area—it seems likely that the current popu-lation is not naturally-occurring(USFWS 2009). Still, Griffith Park iswithin the historical range of the spe-cies and all plants occur in a low-elevation, arid chaparral communitywith a species composition very simi-lar to known natural occurrences.

Another difficult-to-assess rareplant in the park is the San GabrielMountains leather oak (Quercusdurata var. gabrielensis, CaliforniaRare Plant Rank 4.2), which isknown from a single collection in1991. However, more study and col-lection of various scrub oaks in thepark are needed, particularly sinceGriffith Park may lie within an in-trogression zone between the floraof the Santa Monica and San GabrielMountains. This variety appears tobe otherwise unknown in the SantaMonica Mountains.

A map showing the distributionof the above rare species in the parkis provided in Figure 2 on page 20.

LOCALLY RARE SPECIES

Several locally rare species werefound in substantial populations inGriffith Park. Both Nevin’s brickle-bush (Brickellia nevinii) and Ft.Tejon milk-aster (Stephanomeriacichoriaceae) are frequent on rockoutcrops and roadcuts throughoutthe park and even in surroundingresidential areas to the south. East-wood’s manzanita (Arctostaphylosglandulosa ssp. mollis), chaparral pea(Pickeringia montana), and interiorlive oak (Quercus wislizenii var. fru-tescens) occur high atop Cahuengaand Burbank peaks within a densechamise-manzanita chaparral abovethe 1,600-foot elevation level, withsmaller numbers of manzanita and asingle clump of interior live oak nearMt. Hollywood.

“Moss gardens,” where soil hascollected at seeps on rock faces, sup-port a variety of bryophytes, ferns,and rare annuals at Royce Canyonand east along the north side of Mt.Bell, including California saxifrage(Saxifraga californica), Peninsularonion (Allium peninsulare), Fremontstar-lily (Zigadenus fremontii), and(Royce Canyon only) a white-flow-ered form of Cleveland’s shooting-star (Dodecatheon clevelandii). A fewareas of private land adjacent to thepark were found to be rich in scarce

flora, and more probably await dis-covery. The only known popula-tion of dwarf brodiaea (Brodiaeaterrestris var. kernensis) in the east-ern Santa Monica Mountains (A.Gibson, pers. comm.) was also dis-covered by a volunteer during thesurvey.

Uncommon species reaching alocal distributional limit near thepark include erect goldenaster (He-terotheca sessiliflora var. fastigiata),the latifolium subspecies of the wide-spread California fuchsia (Epilobiumcanum), and valley cholla (Cylindro-puntia californica var. parkeri). A fewcommon wildflowers in the parkshow more of a San Gabriel Moun-tains affinity, including Canterburybells (Phacelia minor), one of themost abundant spring annuals inthe park, becoming much more lo-calized farther west in the SantaMonica Mountains.

The population of southern Cali-fornia black walnut (Juglans califor-nica) in the park is so large that itwas not mapped during the survey.Black walnuts form a continuouswoodland/high shrubland on sedi-mentary soils in the southeasterncorner of the park, and this is also adominant species in the chaparralthat cloaks the northern slope of thepark.

PATTERNS OF DIVERSITY

Although areas at the lowest el-evations at the perimeter of the parktended to support fewer target spe-cies, several featured a handful oflocally rare species. Important popu-lations, including the only knownlocal occurrences, were found atmultiple and scattered locations,making it difficult to declare a givenarea unimportant for any rare plant.Not surprisingly, the fewest targetspecies overall were found in thenortheastern corner of the park. Thisarea that has probably seen the mostintensive human-caused impactsover the years, including the con-struction of a large landfill in the

Dwarf broadiaea, a locally rare plant other-wise unknown in the eastern Santa MonicaMountains, was discovered just outside thepark boundary in 2010.


1950s, the installation of a vast net-work of metal irrigation pipe de-signed for fire control (but neverfunctional), and widespread plant-ing of eucalyptus and pines, ostensi-bly for “beautification” starting inthe early 1900s.

It also became clear in our surveythat most scrub-covered slopes in thepark were largely devoid of rareplants, with diversity mainly confinedto microhabitats such as seeps in deepcanyons, sandy/gravelly ridges, mossgardens, rock outcrops, and patchesof grassland on wet clay soils. Areasof the park that combined these fea-tures predictably supported both ahigh diversity of rare species and largepopulations of them.

The identification and documen-tation of these microhabitats arecritical to making management de-cisions that benefit biodiversity, and

the distribution of rare and local-ized plants can be an important firststep in a site’s conservation. The in-volvement of local volunteers notonly contributes information to thisprocess, it helps ensure that the hu-man community will be invested inresource conservation in “Holly-wood’s backyard.”

REFERENCES

Brusha, R.F., ed. 2003. Native and ex-otic flora of Griffith Park: Source: BillEckert. Unpublished checklist.

California Native Plant Society. 2010.Inventory of Rare and EndangeredPlants. Vols. 7–10. http://cnps.site.aplus.net/cgi-bin/inv/inventory.cgi/Home.

Cartifact, Inc. 2007. Griffith Park(map). City of Los Angeles, Officeof Councilman Tom LaBonge.

Consortium of California Herbaria

Nevin’s barberry, one of the rarest plantsin California (in the wild), thrives inGriffith Park, although the entire popula-tion is suspected to have been introduceddecades ago. It is a large shrub found verylocally on loose, gravelly soils withinchaparral, and is endemic to southernCalifornia. Photographs by Gerry Hans.


(CCH). 2011. Data provided by theparticipants of the Consortium ofCalifornia Herbaria. http://ucjeps.berkeley.edu/consortium/.

Cooper, D.S. 2010. Griffith Park RarePlant Survey. October 10, 2010. Un-published report. http://www.friendsofgriffithpark.org.

Cooper, D.S. Forthcoming. Flora ofGriffith Park. Annotated checklist.

Cooper, D.S. and P. Mathewson. 2009.Griffith Park wildlife managementplan. Report submitted to the LosAngeles Department of Recreationand Parks by Cooper EcologicalMonitoring, Inc. http://www.griffithparkwildlife.org.

Gibson, A. and B.A. Prigge. 2003. Re-vised flora of the Santa MonicaMountains. Final Report. Herbar-ium, University of California, LosAngeles.

McAuley, M. 1985. Wildflowers of theSanta Monica Mountains. CanyonPublishing, Canoga Park, CA.

National Park Service (no date). Plantchecklist for the Santa MonicaMountains National Recreation Area.http://www.nps.gov/samo/naturescience/plants.htm.

United States Fish and Wildlife Service.2009. 5-year review for Berberisnevinii (Nevin’s barberry). CarlsbadFish and Wildlife Office. August 14,2009.

Wishner, C. 1997. Flora of the SantaMonica Mountains: Synonymisedchecklist and index. Crossosoma23(1):1-52.

Daniel S. Cooper, Cooper Ecological Moni-toring, Inc., 255 Satinwood Avenue, OakPark, CA 91377, [email protected]

ABOVE: “Moss gardens,” or patches of moist soil within rock outcrops, are key areas for rareplants at Griffith Park. • LEFT: The prickly phlox (Leptodactylon californicum) occurs asscattered plants within the park, mainly in small openings in arid scrub on eroding ridgesand slopes. Its bright pink flowers are unmistakable in spring. • BOTTOM LEFT: The strikingCatalina mariposa lily (Calochortus catalinae) is one of three species of Calochortus in thepark, and is found amid grasses on patches of heavy clay soil, especially those that staymoist through the spring.


A PLEA TO PROTECT WALKER RIDGEby Stephen W. Edwards

(Editor’s Note: The following articleis an abbreviated version of a longerarticle that appeared in The FourSeasons, Journal of the Regional ParksBotanic Garden, Volume 14, Number1, October 2011.)

alker Ridge, on theboundary of Lake andColusa counties be-tween Clear Lake and

the 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 andtimely action of conservation orga-nizations— displays in good yearsby far the best remaining panora-mas of Northern California’s fieldwildflowers. No other area comesclose.

Sediments on the valley floor arestrongly influenced by the magne-sium-rich, calcium-poor ultramaficrocks of Walker Ridge, and this al-most certainly contributes to thewildflower displays. The ridge andthe valley have been a combined des-tination for wildflower lovers, re-searchers, and university class studyfor decades. Visiting the one almostalways entails visiting the other be-cause the contrast between theirnatural histories is fascinating. Thevalley with its vast, flowery panoramais made all the more enchanting bythe bold backdrop of Walker Ridge.

OF NATIONAL PARKQUALITY

Walker Ridge and Bear Valleytogether are clearly of national parkquality. The Carrizo Plain NationalMonument, which includes the plainas well as parts of the Temblor andCaliente ranges in Southern Califor-nia, is an excellent analogy. It is a

park celebrated principally as a rem-nant of the wildflowers of the SanJoaquin Valley as Thomas JeffersonMayfield and John Muir describedthem (1850s and 1860s respec-tively). So it is with Bear Valley andWalker Ridge, which are to the origi-nal, unspoiled Sacramento Valley ofthe early 1800s what the CarrizoPlain National Monument is to theSan Joaquin Valley.

That is why I have long felt thatWalker Ridge—most of which isunder the jurisdiction of the Bureauof Land Management (BLM)—andparts of Bear Valley that might oneday be in public ownership, shouldbecome a national monument.Walker Ridge preserves much of oneof California’s three greatest serpen-tine terrains (the others are The Ce-dars in Sonoma County and the Trin-

WMap showing the proximity of Walker Ridge to Bear Valley. Source: Gregory Gallagher.

Green’s butterweed (Senecio greenei), an orange-hued serpentine endemic found on WalkerRidge, bears some of the most striking flower heads of any sunflower. It has stubbornlyresisted all attempts by horticulturists to propagate it. Also pictured is common woollysunflower (Eriophyllum lanatum). Photograph by Craig Thomsen.


ity ultramafic sheet around Mt. Eddyin Siskiyou County), and these arethree of the greatest serpentine ar-eas on earth. Walker Ridge is a liv-ing classroom of inner coast rangenatural history, from which one cangaze eastward at Sierran snows andthe Pleistocene volcanoes of theSutter Buttes, westward to the ClearLake volcanic field, and north to thesnow-capped peaks of MendocinoNational Forest.

It makes sense to imagine WalkerRidge and parts of Bear Valley asCalifornia Serpentine NationaIMonument. Serpentinite, which,owing to its fortunate chemistry, ac-counts for a greatly disproportion-ate amount of high-quality Califor-nia wildflower habitat, has not re-ceived adequate recognition at a fed-eral (or state) level. Walker Ridgeoffers an opportunity to rectify thatinsufficiency and, at the same time,provide a park of surpassing beautyand fabulous biodiversity that couldbe enjoyed by all.

IMPENDING DEVELOPMENTTHREAT

At the time of this writing, plansare underway by a Canadian energycompany to develop an extensiveand elaborate array of giant wind

turbines, with all their associatedequipment, facilities, pads, yards,and roads, along the serpentine sum-mit of Walker Ridge. Those familiarwith the biology and scenery ofWalker Ridge know that it is the lastplace in California that should beconsidered for any kind of intrusivedevelopment, and that is not merelyowing to its richness. Accessibilityis another critical factor. Here onecan easily drive to many spectacularhabitats, and the whole area is onlyone to two hours by car from majorpopulation centers and universities.

UNPARALLELED RICHNESS,BEAUTY

It has long been known that ser-pentine meadows on Walker Ridgeare the place to go for great displaysof flowers after Bear Valley haslargely dried up. The ridge also of-

fers many open serpentine screeslopes decorated with striking flow-ering gems that cannot endure com-petition on less severe sites. (Screeis a gravely deposit that forms onhillsides where bedrock is weather-ing not far below the surface.) Manyof these plants, too, can be enjoyedin flower in summer. Road banksthemselves, and natural benchesabove them, are full of gorgeousplants. Many of these plants may bedifficult for anyone with mobilityissues to find or reach away fromthe road. Serpentine chaparral alsocontains in its understory and rockyopenings a host of exciting plants.Streams that cut serpentine gorgesalong and near Bartlett Springs Roadcascade over travertine benches and

ABOVE: Serpentine scree slopes along the crest of Walker Ridge are precious preserves forrare, unusual, and beautiful plants. The proposal for a wind farm on the ridge includes anarray of giant wind turbines, along with attendant roads, trenches, etc. Photograph by PhilVan 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 thefloor of Bear Valley, with Walker Ridge in the background, April 1995. Photograph byStephen W. Edwards.

People make pilgrimages to Walker Ridgeto see the rare serpentine milkweed (Ascle-pias solanoana). It can be found only onsliding scree, through diligent searching.Its low, ground-hugging habit makes itintolerant of competition. Photograph byStephen W. Edwards.


bring clear, cool water to lush andextraordinarily diverse riparian veg-etation that includes other, equallybeguiling wildflowers. The botani-cal richness of Walker Ridge is toogreat to enumerate. There are sev-eral running plant lists, and they allkeep growing as new discoveries aremade.

Walker Ridge is not comprisedentirely of serpentinite. Much of thesouthern half north of Highway 20is on Great Valley Sequence sand-stones. These support a more com-mon, less diverse chaparral andwoodland. Nevertheless, the hugefire of 2008 burned much of thispart of the ridge, creating the condi-tions for sheets of wildflowers, andexposing all kinds of lovely plants

that botanists had driven by for de-cades.

Walker Ridge is one of the great-est places left in California. Is thiswhere we would choose to build aconglomeration of huge (nearly 400feet tall) wind towers? Would wescrape off the serpentinite and rareplant jewels for yards, pads, trenches,and tarmac? Do we want to widenWalker Ridge Road for oversizedtrucks, grade away its rich roadbanks, replace tranquility and natu-ral beauty with fluster and rush,oversight, devastation, and weeds?

Walker Ridge is rich in wild-life—foothill yellow-legged frogsseem to inhabit every serpentinestream—and there is important ar-chaeological evidence of native

Americans. Tuleyome, a conserva-tion organization centered in Wood-land, has been working hard to pre-serve Walker Ridge as part of theproposed Berryessa-Snow MountainNational Conservation Area. At thesame time, the California NativePlant Society has formally proposedthat the BLM make the ridge an Areaof Critical Environmental Concern.I am proposing national monumentstatus, but, whatever the name, thecritical thing is that the dignity ofWalker Ridge be recognized withthe honor it deserves.

Stephen W. Edwards, Regional ParksBotanic Garden, c/o Tilden Regional Park,Berkeley, CA 94708-2396, [email protected]


SERPENTINE SOILS AND WHY THEY LIMITPLANT SURVIVAL AND GROWTH

by Earl B. Alexander

erpentine evolves from pris-tine ultramafic rocks formedbeneath the oceans. Rockswhich contain high levels of

iron and very high levels of magne-sium are alien on the continents,where there is more calcium in therocks than magnesium.

Serpentine soils are those thatdevelop from the weathering anddisintegration of ultramafic rocks.They inherit much of their mineralcomposition and chemistry fromthose rocks. Serpentine soils are sodifferent from others that relativelyfew plants are adapted for survivalin them. Ultramafic rocks and soilscover only about 1% of the land areain California, yet 12% of the en-demic species in the state are re-stricted to serpentine soils (Saffordet al. 2005). How are ultramaficrocks and serpentine soils differentfrom non-ultramafic ones, and whyare serpentine soils so difficult forplants to cope with?

ROCKS FROM WHICHSERPENTINE SOILSDEVELOP

Serpentine is ecological jargonfor ultramafic rock. The commonultramafic rocks in California areperidotite and serpentinite. Peridot-ite is the ultramafic rock producedby crystallization of magma beneaththe oceans, and serpentinite is pro-duced by the hydrothermal alter-ation of peridotite. Serpentine wasdesignated the state rock in Califor-nia before geologists decided to callthe rock serpentinite and restrict ser-pentine to the name of the dominantmineral in serpentinite (Wagner1991).

The story of ultramafic rocks andserpentine soils begins in the mantle.The mantle is between the core andthe crust of Earth. Geologists be-lieve that the composition of theupper mantle is comparable to thatof lherzolite, which is a type of peri-dotite. Below the oceans, partialmelting in the upper mantle pro-duces a magma that has more of theelements with larger cations (posi-tive ions), such as potassium andcalcium, and less of the elementswith smaller cations, such as mag-nesium, than in lherzolite.

Much of the magma risesthrough the ocean crust at spread-ing centers, or ridges, and flows out-ward under seawater, cooling rap-idly to form rounded blobs of lavathat become a distinctive feature ofpillow basalt. Magma that does notrise to the top of the ocean crustcools more slowly and forms gab-bro, an intrusive igneous rock thatis the equivalent of basalt, anextrusive igneous rock. Beneath thegabbro, residue from partial meltingin the upper mantle forms harz-

burgite, which is another type ofperidotite. It is in harzburgite thatthe elements with smaller cationssuch as magnesium, chromium,manganese, iron, cobalt, and nickelare concentrated following partialmelting of lherzolite. The sequenceof rocks from basalt, through aswarm of dikes that have fed basaltupward, and gabbro to harzburgite,is called ophiolite.

Ophiolite is produced near thecenters of large blocks of ocean crustcalled plates. The ocean plates moveaway from the spreading centers to-ward continental plates. Along the

S

ELEMENTABBREVIATIONS

C carbon

Ca calcium

Co cobalt

Cr chromium

H hydrogen

K potassium

Mg magnesium

N nitrogen

Ni nickel

O oxygen

P phosphorous

S sulfur

Ophiolite, which consists of basalt, feederdikes, gabbro, and harzburgite (peridotite).Serpentine forms by alteration of theminerals (olivine and pyroxenes) inperidotite.


way, the ophiolite cools, and sedi-ments accumulate on the oceanfloor. As the ophiolite cools, it be-comes heavier. By the time spread-ing ocean crust reaches a continent,it is generally cool and heavy enoughto sink under the edge of the conti-nental plate into the mantle, a pro-cess called subduction.

Subducted plates sink for manymillions of years, and some ophiolitemay reach depths of about 410 kilo-meters near the bottom of the uppermantle. Sediments are commonlyscraped off of the plates and accu-mulate in long trenches where theplates sink beneath continental mar-gins, with some sediments beingdragged under continental marginsand plastered against the continen-tal crust, underplating it. Plates thathave lost their sedimentary cover

sink to depths where they begin tomelt. Partial melting of the platesand adjacent continental crust pro-duces magmas that may rise to thesurface and form volcanoes. Suchvolcanoes that skirt the Pacific Oceancomprise a volcanic chain aptlycalled the ring of fire. The CascadeMountain Range from Lassen Peakin Northern California to MountBaker in Washington is a link in thischain of volcanoes.

The ultramafic rocks in sub-ducted ophiolites must reach theground surface to produce serpen-tine soils. Rocks can be raised to thesurface by faulting, or the rocks canbe folded and rocks in the folds ex-posed by erosion of the overlyingrocks. The California Coast Rangesare comprised largely of the Fran-ciscan complex of accreted oceanic

terranes, and ophiolites have beenthrust over them. Granitic bodiescalled plutons, and volcanic depos-its are commonly added to accretedterranes after the terranes are at-tached to a continent. The SierraNevada batholith is an amalgamationof many plutons, for example, andvolcanic rocks in the Coast Ranges,such as those of Mount St. Helens,postdate accretion of the Franciscancomplex.

Most peridotite (mainly harz-burgite) that we see in Californiahas been at least partially altered toserpentinite. The alteration, a pro-cess called serpentinization, beginsunder the ocean and continues evenafter harzburgite has been incorpo-rated into the continental crust.Serpentinization is a low tempera-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 onmoderately deep serpentine soils on the right, in contrast to a dense mixed conifer forest on nonserpentine soils on the left. Allphotographs by the author.


water is added to the olivine andpyroxenes in harzburgite, and somecalcium and silicon are lost frompyroxenes:

harzburgite + water ===>serpentinite + calcium hydroxide +

silicaThe water that reacts with harz-

burgite generally lacks carbon diox-ide; its main role is to provide oxy-gen and hydroxyl ions used in theformation of serpentine, which con-tains more of these than the miner-als in harzburgite. Water left overfrom the serpentinization processcontains calcium and is alkaline (pH> 11). It is responsible for convert-ing some of the gabbro and basalt inophiolites to calcium silicate (rodin-gite). Water from the serpentiniza-tion of harzburgite that flows fromthe rocks to the ground surface re-acts with bicarbonates in meteoricwater (water that has gained carbondioxide in the atmosphere or soils)to precipitate the calcium and formtravertine. Note that carbon dioxidedissolved in water reacts with it toform carbonic acid that dissociatesinto hydrogen (H+) and bicarbonate(H(CO

3)-) ions:

Ca(OH)2 + H+, H(CO

3)- ===> Ca

(CO3) + 2H

2O

Ca-hydroxide + carbonic acid ===>travertine + water

PLANT GROWTH ONSERPENTINE SOILS

Serpentine soils are those thathave peridotite or serpentinite par-ent materials. These two parent ma-terials commonly produce soils withdifferent physical properties, but theplants appear to respond more tothe chemical properties that are simi-lar for soils developed from eitherparent material. Serpentine soilshave a broad range of depths andwater-holding capacities, as do soilswith most other kinds of parent ma-terials (Alexander et al. 2007).Therefore, we can concentrate onthe chemistry to learn why plantsrespond differently inserpentine soils thanin other soils.

The main ele-ments required byplants are H, C, N, O,Mg, P, S, K, and Ca.The first four of theseelements are in at-mospheric gases, orwater, from whichplants obtain them ei-ther directly or indi-rectly. Elements forwhich plants are de-pendent on the soilparent materials aremainly Ca, Mg, K,and P. Because Mg

concentrations are very high and Caconcentrations are very low in ser-pentine soil parent materials, the Ca/Mg ratios are very low compared tothose in average continental crustand soils.

The very high Mg concentra-tions, relative to Ca, limit plant up-take of Ca from soils. High Mg (andlow Ca) concentrations in plants in-terfere with metabolic processes thatrequire Ca, excluding plants thathave not developed mechanisms todeal with these Ca/Mg imbalances.If supplemental Ca is added to aserpentine soil, P becomes a plantgrowth-limiting element. Plantscommonly respond favorably to the

LEFT: Travertine draped over the rocks and forming a small pool where alkaline water produced from the serpentinization of peridotiteemerges from the ground, Sonoma County. • MIDDLE: Pillow basalt downstream from the outlet to Nicasio Reservoir, Marin County. • RIGHT:Serpentinization in an exhumed rock.

A pictorial view of serpentinization. Water has entered alongfractures in peridotite and caused serpentinization, whichdiminishes from the fractures toward cores that are onlyslightly serpentinized peridotite.


AID TO TERMINOLOGYaccreted – added to a continent

cations – positively charged ions

gabbro – intrusive (produced below ground) equivalent of extrusivebasalt

harzburgite – ultramafic rock produced from lherzolite (definedbelow) beneath ocean crust

hyperaccumulators – plants that accumulate more than 0.1% byweight of nonessential elements (e.g., Ni or Co)

lherzolite – ultramafic rock believed to reflect the composition of theupper mantle, lherzolite yields plutonic (intrusive) harzburgite,plus intrusive gabbro and extrusive basalt

mafic lava – molten rock, or magma, that flows over the surface of theearth and cools to form basalt, an aluminum silicate rock withlarge concentrations of alkaline earth cations (Ca and Mg) and iron

ophiolite – a section of the Earth’s oceanic crust and upper mantlethat has been uplifted and exposed above sea level

peridotite – ultramafic rock formed by crystalli1zation of magmaunder the ocean

pillow basalt – mafic lava that flows over the seafloor and cools tosolidify in large blobs resembling pillows

pluton – a body of rock (see plutonic rock) formed at depth

plutonic rock – rock that crystallizes slowly from magma at greatdepths to form crystals visible without magnification

pyroxenes – magnesium silicate minerals, lacking aluminum

serpentine – dominant mineral in serpentinite (and previously thename of the rock)

serpentinite – a greenish metamorphic rock consisting largely ofserpentine

subduction – sinking or pushing of an oceanic plate beneath anotherplate

terrane – alien crustal block that has an origin different from that ofadjacent rocks

trichomes – hairlike appendages to plant epidermal cells

ultramafic rocks – rocks consisting mainly of magnesium silicateminerals, with much iron substituting for some of the magnesium

vacuoles – spaces within cells that are filled with liquids

addition of N to serpentine soils,but the same is also true for plantsgrowing in other kinds of soils.

The elements that are muchmore concentrated in serpentinesoils than in others may be toxic toplants. These elements are Cr, Ni,and Co. Some plants such as Sargentcypress (Cupressus sargentii) andleatheroak (Quercus durata) have de-veloped mechanisms to deal with

elevated concentrations of these el-ements. They may selectively ex-clude the elements from root up-take, confine them to roots, or accu-mulate them as chelated forms orprecipitates in the vacuoles of epi-dermal cells in leaves or concentratethem in trichomes on the leaves.

Plants that accumulate hundredsof times more of an element thanmost other plants are called hyper-

accumulators. There are hundredsof plant species that hyperaccumu-late Ni, and a few that hyperaccu-mulate Co. There are only two Nihyperaccumulating species in Cali-fornia, and both are in the mustard(Brassicaceae) family: milkwortjewel flower (Streptanthus polyga-loides) and two varieties of alpinepennycress (Noccaea montana)(Reeves et al. 1983). The only Cohyperaccumulating species arefound in Africa.

Plants that thrive on serpentinesoils have unusual capabilities to uti-lize calcium when the alkaline earthelements are dominated by magne-sium, and to tolerate concentrationsof cobalt and nickel that are toxicto most plants. Plants of intertidalzones are also adapted to grow insoils with low Ca/Mg ratios, but theydo not have to cope with potentiallytoxic concentrations of cobalt andnickel. Although relatively fewplants are adapted to grow in ser-pentine soils, many of them can-not compete with other plants onnonserpentine soils and thus areendemics confined to serpentinesoils.

REFERENCES

Alexander, E.B., R.G. Coleman, T.Keeler-Wolf, and S. Harrison. 2007.Serpentine Geoecology of WesternNorth America. Oxford UniversityPress, New York.

Reeves, R.D., R.M. Macfarlane, andR.R. Brooks. 1983. Accumulation ofnickel and zinc in western NorthAmerican genera containing serpen-tine-tolorant species. American Jour-nal of Botany 70: 1297-1303.

Safford, H.D, J.H. Viers, and S.P.Harrison. 2005. Serpentine ende-mism in the California flora: A data-base of serpentine affinity. Madroño52: 222-257.

Wagner, D.L. 1991. The state rock ofCalifornia: Serpentine or serpenti-nite? California Geology 44: 164.

Earl Alexander, Soils and Geoecology,1714 Kasba Street, Concord, CA 94518,[email protected]


SERPENTINE ENDEMISM IN THE CALIFORNIA FLORAby Hugh D. Safford

ltramafic “serpentine” soilsand rock outcrops supportan exceptionally high num-ber of endemic plant taxa

in California, far out of proportionto their limited representation onthe California landscape. California“endemic” plant taxa are species,subspecies, and varieties that are re-

stricted in their range to California;ultramafic endemics are only foundon ultramafic soils. About 11% ofthe taxa found on the California RarePlant Rank 1B (formerly CNPS List1B, critically rare taxa) are eitherstrictly or broadly endemic to ultra-mafic substrates. Rare native taxathat are not serpentine endemics alsofind important habitat footholds onultramafic substrates, where they canescape competition from more ag-gressive, often non-native species.

WHAT ARE SERPENTINESUBSTRATES?

By definition, ultramafic rocksare igneous and metamorphic rocksthat contain less than 45% by weightsilica (SiO

2) or, alternatively, sup-

port a mineral content that is greaterthan 90% “mafic” (i.e., magnesiumand iron silicate minerals like oliv-ine, pyroxenes, or hornblendes)(Ehlers and Blatt 1982). Ultramaficrocks originate primarily in theearth’s mantle, and find their way tothe surface by way of plate tectonicsand mountain building, and subse-quent erosion of overlying geologiclayers.

Ultramafic rocks and the soilsthat develop on them (often collo-quially called “serpentine” after theirmost important mineral constitu-ents) are characterized by criticallylow levels of most principal plantnutrients (nitrogen, phosphorus,potassium, calcium), exceptionallyhigh levels of magnesium and iron,and a suite of highly toxic trace ele-ments including chromium, nickel,and cobalt (Alexander et al. 2007).In all, ultramafic rocks occupy an

area of approximately 2,300 squaremiles in California (Harrison et al.2000), which makes up around 1.5%of the state’s area. However, as EarlAlexander notes in the accompany-ing article, inexactitude in mappingmay inflate that area somewhat.

NUMBERS AND TYPES OFSERPENTINE PLANTS

Serpentine endemism is a keycomponent of the diversity of theCalifornia flora (Raven and Axelrod1978, Kruckeberg 2002, Safford etal. 2005). In 2005, we carried out anassessment of serpentine endemismin the California flora (Safford et al.2005). We found that of the 1,416endemic species listed in The JepsonManual, 1st edition (Hickman 1993),about 176 could be ranked as either“strict” or “broad” serpentine en-demics (“strict” having over 95% ofits occurrences on serpentine soilsand “broad” having over 85%). Thisconstituted about 12.7% of the Cali-fornia endemic flora.

Since the 2005 analysis, manychanges have occurred in the tax-onomy and classification of the Cali-fornia flora, and a surprising num-ber of new species have been de-scribed. Information from the Con-sortium of California Herbaria(CCH, http://ucjeps.berkeley.edu/con-sortium/) records, The Jepson Manual,2nd edition (Baldwin et al. 2012),and the scientific literature was usedto decide how to rank newly de-scribed, combined, or split taxa withrespect to their preference for ser-pentine soils.

This new information shows thatapproximately 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 talusfritillary (Fritillaria falcata). All are restricted to a handful of outcrops in the North orSouth Coast ranges. Photographs by Rick York, Bob Case, and John Game.

U


TABLE 1. NUMBERS OF SERPENTINE ENDEMIC ANDNEAR ENDEMIC TAXA FOR PLANT FAMILIES WITH ATLEAST THREE ENDEMIC TAXA ON ULTRAMAFICSUBSTRATES IN CALIFORNIA

FAMILY SERPENTINE AFFINITY SCORE

Strict Broad Near Totalendemics endemics endemics taxa1

Asteraceae 26 11 8 105

Brassicaceae 19 3 4 37

Polygonaceae 11 8 3 41

Liliaceae 6 6 3 37

Apiaceae 7 3 3 32

Orobanchaceae 7 3 0 19

Linaceae 8 1 0 14

Ericaceae 5 3 2 15

Alliaceae 4 3 4 24

Fabaceae 4 3 3 24

Polemoniaceae 6 1 1 18

Boraginaceae 6 0 5 24

Caryophyllaceae 5 1 3 18

Lamiaceae 4 2 3 17

Campanulaceae 3 2 3 13

Crassulaceae 5 0 2 13

Cyperaceae 2 3 2 11

Onagraceae 3 2 2 13

Rhamnaceae 4 1 1 15

Agavaceae 3 1 0 6

Convolvulaceae 1 3 1 6

Rubiaceae 3 1 0 8

Themidaceae 2 2 3 10

Montiaceae 0 3 2 16

Plantaginaceae 1 2 2 15

Poaceae 1 2 0 18

Totals (all families) 164 79 72 661

Familes are ordered by their relative numbers of serpentine endemics (strict plusbroad endemics). Strict endemics have >95% of their documented occurrenceson ultramafic substrates; broad endemics (>85%-94.9%); near endemics (>75%-84.9%). The latter group is the transitional group from “strong indicators” tobroad endemics in Safford et al. (2005).

1Total of all taxa in our database, from strict endemics to indifferent taxa.

to ultramafic substrates in Califor-nia, including 176 full species. Thesenumbers are very close to our 2005numbers (246 taxa, 176 full spe-cies), although some of the nameshave changed, some taxa weredropped from the list, and some newtaxa were added. In addition, due tochanges in taxonomy and two de-cades of additional field collection,the estimated number of plant spe-cies endemic to California has actu-ally dropped to 1,315 (Baldwin etal. 2012), so the contribution of ser-pentine endemism to the Californiaendemic flora has risen to 13.4%!Table 1 provides a summary of ser-pentine endemic richness in Cali-fornia. Altogether, the families inTable 1 contribute almost 90% (216of 243) of the total serpentine en-demic diversity in California.

Asteraceae is the top family inall categories listed, while secondand third place are held by Brassi-caceae and Polygonaceae. Figure 1plots the families listed in Table 1.The first eight families include morethan 50% of the taxa in my endemicdatabase. Asteraceae includes about15% of the serpentine endemic taxain the list, a similar proportion to itscontribution to the total Californiaendemic flora. Five of the next sixfamilies (all but Orobanchaceae) aresubstantially overrepresented in theserpentine flora, which means theyhave been important evolutionarysources of new species on serpen-tine soils.

The most diverse genera of ser-pentine endemism are listed in Table2, topped by Eriogonum and Streptan-thus, and followed by Hesperolinonand Arctostaphylos. The genera inTable 2 include more than 50%of the total taxa endemic to serpen-tine soils in California. Basically,most of the serpentine endemic plantdiversity in California is driven byonly a few dozen genera (in only afew dozen families) that have evolvedmany species on serpentine (andsometimes other harsh) substrates.

Probably the flagship for serpen-


tine endemic plants inCalifornia is the genusStreptanthus (Brassica-ceae), which includes atleast 16 plant species orsubspecies that are strictlyor broadly endemic to ser-pentine soils. Streptanthus(commonly called the“jewelflowers”) is an ex-ample of a neoendemictaxon, a relatively recentlyevolved plant group thatcontinues to actively gen-erate new species. Ex-amples include Socratesmine jewelflower (Strep-tanthus brachiatus), whichis restricted to a few ser-pentine locations at the junction ofLake, Napa, and Sonoma counties,and milkwort jewelflower (Strep-tanthus polygaloides), a species fromthe Sierra Nevada which has evolvedthe rare ability to hyperac-cumulatenickel, a highly toxic heavy metalthat is usually deadly in the concen-trations found in the plant.

Of the 243 serpentine endemicsidentified in the updated assessment,86% are forbs (210 out of 243), withone-third of those being annuals andtwo-thirds perennials (Figure 2).Some of the more charismatic forbsare in the old lily family, now splitinto numerous families. Examplespictured are Indian Valley brodiaea(Brodiaea rosea), San Luis mariposalily (Calochortus obispoensis), andtalus fritillary (Fritillaria falcata), allof which are restricted to a handfulof ultramafic outcrops in the Northor South Coast Ranges.

Graminoids (grasses, sedges, andrushes), shrubs, and trees compriseonly 14% of the endemic serpentineflora (Fig. 2). Leather oak (Quercusdurata var. durata) and musk brush(Ceanothus jepsonii) are serpentineendemic shrubs that often grow to-gether in the North Coast Ranges.One of the few tree taxa that is aserpentine endemic is Sargent cy-press (Hesperocyparis sargentii),which is found from Mendocino

TABLE 2. GENERA WITH AT LEAST FOUR SERPENTINEENDEMIC TAXA IN CALIFORNIA

Genus Family Endemic taxa

Eriogonum Polygonaceae 16

Streptanthus Brassicaceae 16

Hesperolinon Linaceae 9

Arctostaphylos Ericaceae 8

Allium Alliaceae 7

Lomatium Apiaceae 7

Packera Asteraceae 6

Calochortus Liliaceae 5

Carex Cyperaceae 5

Cordylanthus Orobanchaceae 5

Calystegia Convolvulaceae 4

Castilleja Orobanchaceae 4

Cirsium Asteraceae 4

Erigeron Asteraceae 4

Fritillaria Liliaceae 4

Galium Rubiaceae 4

Harmonia Asteraceae 4

Lessingia Asteraceae 4

Minuartia Caryophyllaceae 4

Monardella Lamiaceae 4

Phacelia Boraginaceae 4

FIGURE 1. CALIFORNIA PLANT FAMILIES WITH AT LEAST THREE SERPENTINE ENDEMICTAXA, AS A PERCENTAGE OF THE TOTAL CALIFORNIA SERPENTINE ENDEMIC FLORA


County to Santa Barbara County,mostly at sites with moister soils orperiodic summer fog.

Since most ultramafic outcropsare very limited in size, many ser-pentine endemic plant taxa also havesmall geographic ranges. Combinedwith the occurrence of many ser-pentine outcrops in populated (andgrowing) coastal and foothill urbanand suburban areas, this has led toan overrepresentation of serpentineendemic plants among the state’sendangered, rare, and sensitive planttaxa (CNPS Inventory of Rare andEndangered Plants, http://www.rareplants.cnps.org/).

The conservation status of thetaxa in my database has not yet beenupdated, but in 2005, 186 (77%) ofthe taxa in the endemic category(strict plus broad endemics) wereincluded in the CNPS Inventory ofRare and Endangered Plants (Saffordet al. 2005). California Rare PlantRank 1B, which includes federaland state threatened and endan-gered taxa, as well as taxa that CNPSbelieves warrant listing due to theirrarity, included 105 serpentine en-demics.

Serpentine endemics listed asfederal or California endangeredinclude Tiburon Indian paintbrush(Castilleja affinis ssp. neglecta),which is restricted to a couple ofultramafic outcrops in the northSan Francisco Bay Area; coyote

ceanothus (Ceanothus ferrisiae),found only in the low mountainssoutheast of San Jose; and Kellogg’sbuckwheat (Eriogonum kelloggii),which grows only at the top of RedMountain in Mendocino County.

WHERE SERPENTINEENDEMICS ARE FOUND

Harrison et al. (2006) carried outa detailed statistical analysis of thefactors influencing the geographicdistribution of serpentine endemismin California. They found that localrichness (the number of serpentineendemic plants) is primarily drivenby an interaction between the re-

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

All three of these serpen-tine endemic species arelisted as federal or Cali-fornia endangered. TOP

TO BOTTOM: Coyote ceano-thus (Ceanothus ferrisiae),Kellogg’s buckwheat (Erio-gonum kelloggii), TiburonIndian paintbrush (Cas-tilleja affinis ssp. neglec-ta). Photographs by NealKramer, Steve Edwards,and Kaylea Eickhoff.

FIGURE 2. LIFEFORMS OF SERPENTINE ENDEMIC PLANT TAXA IN CALIFORNIA


Localities that have rockier soils havehigher richness of endemics, whileoverstory cover, litter cover, and therelative balance of soil magnesiumversus calcium have curvilinear(“hump-shaped”) relationships tolocal endemic richness, such thathigh and low values of these vari-ables support less richness and in-termediate values support higherrichness.

Many, if not most, serpentineendemics are confined to ultrama-fic habitats (Harrison et al. 2006)

precisely because the harsh rockysubstrates allowed them to escapecompetition (Brooks 1987). At thesame time, some ultramafic sitesare too harsh even for serpentineendemics (Harrison et al. 2006). Dueto their low soil nutrient status, ul-tramafic outcrops tend to supportrelatively slow growing vegetationand low levels of plant biomass. InCalifornia, with its Mediterranean-like climate and frequent fires, ul-tramafic outcrops tend to burn lessoften than surrounding, more fer-tile substrates (Safford and Harrison2004). Thus serpentine areas mayprovide refuge from both bioticcompetition and ecological distur-bances like fire (Safford and Mallek2010). The abundance of serotinousconifers like knobcone pine (Pinusattenuata, not an endemic) andMacnab cypress (Hesperocyparis mac-nabiana, a broad endemic) on ser-pentine soils is probably primarilydue to the lower frequencies of firein these locations. (Serotinous co-

nifers are those whose cones onlyopen and release seed at high tem-peratures, usually during fires.)

Figure 3 portrays the distribu-tion of serpentine endemic taxa bygeographic region. The center of ser-pentine endemism in the CaliforniaFloristic Province is in the NorthCoast and Klamath Ranges of north-western California (and southwest-ern Oregon, not included here). Inaddition, most serpentine endemicsare found in habitats below 1500meters (5,000 feet) (Figure 4).

At the regional scale, Harrisonet al. (2006) found that serpentineendemic richness is best explainedby a model including regional pre-cipitation and regional vegetationproductivity, the area of ultramaficrock in the region, and how long theultramafic rocks have been avail-able for plant colonization. Regionswith higher endemic diversity werethose with higher precipitation and(regional) productivity, more ultra-mafic rock, and longer availability

The greatest number of serpentine endemics in California are to be found in the Klamathand North Coast Ranges. ABOVE: Flame ragwort (Packera greenei), one of the few orange-flowered members of the sunflower family. Photograph byDick O’Donnell. • LEFT: Siskiyouinside-out flower (Vancouveria chrysantha), a shrub. Photograph by David McClurg.


of ultramafic substrates for coloni-zation.

This helps to explain the pat-terns in Figs. 3 and 4, as the Kla-math and North Coast Ranges arethe wettest part of California, andsupport the most extensive ultra-mafic outcrops and oldest serpen-tine soils in the state. Examples ofinteresting Klamath and North CoastRange serpentine endemics includeserpentine milkweed (Asclepias sola-noana), surely one of the strangestlooking plants in the California flora;flame ragwort (Packera greenei), oc-casionally found on nonserpentinesoils and one of the few orange-flowered members of the sunflowerfamily; and Siskiyou inside-outflower (Vancouveria chrysantha), ashrub whose strange flowers re-semble the shooting star.

Reduced richness of endemics athigher elevations is due principallyto 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 highelevation serpentine endemics inCalifornia, including Trinity buck-wheat (Eriogonum alpinum), and Mt.Eddy draba (Draba carnosula), bothfound about 2,100 meters (7,000feet) in the Scott Mountains west ofMt. Shasta.

In summary, the “average” Cali-fornia serpentine endemic species isa dicotyledonous, perennial forbfrom a short list of plant familiesand genera, rare enough to be listedas warranting conservation concern,and restricted (primarily) to ultra-mafic soils because the difficult en-vironment filters out faster-growingcompetitors. The highest levels ofendemic richness on ultramafic sub-strates in California are found onrelatively harsh (although not ex-treme) soils in areas of relativelyhigh precipitation and vegetationproductivity, at low to moderate el-evations, in regions where large ar-eas of ultramafic rocks have been

available for long enough periods oftime to provide for the evolution ofa large species pool of serpentine-adapted plant taxa.

Recent research on the effects ofcurrent and projected future climatictrends on narrowly distributed en-demic plants (Damschen et al. 2010)suggests that we should be worriedabout the long-term prospects formany of California’s serpentineendemics. The extreme warming

that is projected for the 21st centuryis especially likely to impact plantswhose habitat is restricted to dis-junct patches of soil scattered acrossthe landscape. It will be expensiveand both logistically and politicallydifficult to conserve serpentine en-demic taxa on a species-by-speciesbasis. Conservation of the remain-ing significant areas of unprotectedserpentine habitat, especially in andaround urban and suburban areas in

FIGURE 3. GEOGRAPHIC DISTRIBUTION OF SERPENTINE ENDEMIC TAXA IN CALIFORNIA

“Total endemics” includes all Califiornia serpentine endemic taxa present in a given region. “Restrictedendemics” includes only those taxa restricted to a given region.

FIGURE 4. ELEVATIONAL DISTRIBUTION OF SERPENTINE ENDEMIC TAXA IN CALIFORNIA,AS WELL AS ALL TAXA IN THE SERPENTINE AFFINITY DATABASE

Taxa are counted in every elevational belt where they occur.


the Bay Area, southernNorth Coast Ranges, andSierra Nevada foothills,would be a good—althoughcertainly not sufficient—alternative.

REFERENCES

Alexander, E.B., R.G.Coleman, T. Keeler-Wolf, and S.Harrison. 2007. Serpentine Geoecol-ogy of Western North America. Ox-ford University Press, New York, NY.

Baldwin, B.G., D.H. Goldman, D.J. Keil,R. Patterson, T.J. Rosatti, and D.H.Wilken, eds. 2012. The JepsonManual: Vascular plants of California.2d edition. University of CaliforniaPress, Berkeley, CA.

Brooks, R.R. 1987. Serpentine and itsVegetation. Dioscorides Press, Port-land, OR.

Damschen, E.I., S. Harrison, and J.B.Grace. 2010. Climate change effectson an endemic-rich flora: Resurvey-

ing Robert H. Whittaker’s Siskiyousites (Oregon, USA). Ecology 91:3609-3619.

Ehlers, E.G., and H. Blatt. 1982. Petrol-ogy. W.H. Freeman and Company,San Francisco, CA.

Grime, J.P. 2001. Plant Strategies, Veg-etation Processes, and EcosystemProperties. 2d ed. Wiley, Chichester,UK.

Harrison, S.P., J.H. Viers, and J.F.Quinn. 2000. Climatic and spatialpatterns of diversity in the serpen-tine plants of California. Diversityand Distributions 6: 153-161.

Harrison, S.P., H.D. Safford, J. Grace,

J.H. Viers, and K.F. Davies. 2006. Re-gional and local species richness inan insular environment: Serpentineplants in California. Ecological Mono-graphs 76: 41-56.

Hickman, J.C., ed. 1993. The JepsonManual: Higher Plants of California.University of California Press, Ber-keley, CA.

Jaffré, T. 1992. Floristic and ecologicaldiversity of the vegetation on ultra-mafic rocks in New Caledonia. InThe Vegetation of Ultramafic (Serpen-tine) Soils: Proceedings of the First In-ternational Conference on SerpentineEcology, eds. Baker, A.J.M., J. Proc-

The genus Streptanthus andgenus Eriogonum top the list ofthose containing the greatestnumber of serpentine endemicspecies in California. Picturedare (CLOCKWISE FROM TOP LEFT):Socrates mine jewelflower(Streptanthus brachiatus), milk-wort jewelflower (Streptanthuspolygaloides), and Trinity buck-wheat (Eriogonum alpinum).Photographs by Dick O’Donnell,Bob Case, and Julie KiersteadNelson.


tor, and R.D. Reeves, 101-107. In-tercept Ltd., Andover, UK.

Kruckeberg, A.R. 1984. CaliforniaSerpentines: Flora, Vegetation, Ge-ology, Soils and Management Prob-lems. University of California Press,Berkeley, CA.

Kruckeberg, A.R. 2002. Geology andPlant Life: The Effects of Landformsand Rock Types on Plants. Universityof Washington Press, Seattle, WA.

Raven, P.H., and D.I. Axelrod. 1978. Ori-gins and relationships of the Califor-nia flora. University of California Pub-lications in Botany, Vol. 72. Universityof California Press, Berkeley, CA.

Reeves, R.D., A.J.M. Baker, A. Borhidi,and R. Berazain. 1999. Nickel hyper-accumulation in the serpentine floraof Cuba. Annals of Botany 83: 29-38.

Safford, H.D., and S. Harrison. 2004.Fire effects on plant diversity in ser-pentine vs. nonserpentine chaparral.Ecology 85: 539-48.

Safford, H.D., and C.R. Mallek. 2010.Disturbance and diversity in low pro-ductivity ecosystems. In Serpentine:

TOP: Mt. Eddy draba (Draba carnosula) isamong the comparatively few serpentineendemics found above 7,000 feet. Photo-graph by Steve Matson. • BOTTOM LEFT:Stebbins’ tarweed (Harmonia stebbinsii) isan endangered species found on shallow,rocky soils, and is on California Rare PlantRank 1B.2 (formerly CNPS List 1B.2).Photograph by Julie Kierstead Nelson.BOTTOM RIGHT: There are three varieties offountain thistle found in and around seepsand streams. All are strict serpentine en-demics. This is Cirsium fontinale var. cam-pylon. Photograph by Bob Case.

The Evolution and Ecology of a ModelSystem, eds. S.P. Harrison and N.Rajakaruna, 249-274. University ofCalifornia Press, Berkeley, CA.

Safford, H.D., J.H. Viers, and S.Harrison. 2005. Serpentine ende-mism in the California flora: A data-base of serpentine affinity. Madroño52: 222-257.

Hugh Safford, USDA Forest Service, Pa-cific Southwest Region, 1323 Club Drive,Vallejo, CA 94592, [email protected]


n 1962 a colleague and I drove tothe Mojave Desert from Sacra-mento for our first botanical col-lecting trip to this area. Sometime

in the middle of the night, as wewere driving downslope into north-ern reaches of the desert, I woke upfrom a deep sleep in the passengerseat to see the strangest lookingplants I had ever seen.

There they were within the pe-ripheral glow of the headlights, alienforms that turned out to be Joshuatrees (Yucca brevifolia), Spanishbayonets (Yucca baccata), and blad-der pods (Isomeris arborea) dottingthe upper elevations of the MojaveDesert. As it was to the earliest ex-plorers when they landed on theshores of North America, this was

the New World to me, full of wildcreations and countless new discov-eries.

Even though nearly 50 yearshave passed and I now feel comfort-ably familiar with many of the plantsof the Mojave and Sonoran Deserts,I still feel the excitement of ventur-ing into the “New World” each timeI enter this unique ecosystem. And

PLANT GALLS: DESERT TREASURESby Ron Russo

The Joshua tree (Yucca brevifolia) is the ubiquitous tree of the Mojave Desert. This specimen is growing in Red Rock Canyon State Parkin the eastern Mojave next to fourwing saltbush (Atriplex canescens). The Mojave is home to many plant galls, including some of the newlydiscovered ones mentioned in this article. All photographs by the author except as noted.

I


with each trip, I encounter new spe-cies that only stir the imagination ofwhat is yet to be discovered. Overthese many years, I have made hun-dreds of pilgrimages to these desertsin search of plant galls—tumor-likegrowths on plants induced by in-vading organisms to house their lar-vae, in the case of insects—and tostudy seasonal changes to the plantsthat host them.

Plant galls are specific in size,shape, and color to each species ofinsect that induces them. Further-more, gall insects have evolvedstrictly dependent on either a genusof plants or a species within. A newspecies of gall midge, for example,was discovered in 2004 on desertholly (Atriplex hymenelytra) that doesnot occur on any other host plant.The exact mechanism(s) responsiblefor species-specific designs in plantgalls and species-specific host selec-tion has yet to be uncovered. Todate, I have found 101 species ofthese gall-inducing insects that aredependent on desert plants duringtheir lifecycles. And of these, 38 werepreviously unknown to science.

Some shrubs support relativelylarge numbers of gall insects. Forexample, at least 16 species of gallmidges are found only on the leavesand flowers of creosote bush (Larreatridentata). The creosote stem gallmidge (Asphondylia auripila) in-duces the largest gall, about the sizeof a golf ball, on these resilientshrubs, and it was ground andsmoked by the Sere Indians of theSouthwest. Finding other species,however, like the galls of the creo-sote antler gall midge (A. digitata)that look like tiny moose antlers, orthe galls of the leaf club gall midge(A. pilosa), takes careful and disci-plined observation.

My next favorite among deserthosts is rabbitbrush (Chrysothamnusspp.), which is host to 15 species ofgall insects, including a few newspecies. One such new species, foundin 2004, induces a cotton-like ter-minal flower gall on rubber rabbit-

brush, (C. nause-osus) in the north-ern Mojave. Later, Ifound the same gallinsect on yellow rab-bitbrush (C. viscidi-florus), closer to ourhome at the time inMount Shasta, whichallowed me to makeweekly visits over aperiod of severalmonths and collectthe galls at just theright moment tosuccessfully rear theadult midges. Thesewere later identifiedas a new species ofmidge in the genus,Rhopalomyia. Rab-bitbrush also playshost to one of therarest and strangest interrelation-ships in the insect world where thegall midge (Rhopalomyia bigelo-viae) induces a gall in the gall of thebubble gall tephritid (Aciurina trixa).These “endogalls” as they are called,represent the ultimate to me inmicro-niche evolution.

Another strange occurrence in-volves Cooper’s boxthorn (Lyciumcooperi). I had been studying a par-ticular small group of plants nearHalloran Pass off Highway 15 eastof the small desert community ofBaker. For three years I had col-lected elliptical stem galls belongingto a new species of moth, trying torear the adults (later identified asGnorimoschema sp.). During thesethree years, the stem galls were theonly galls found on the specificshrubs being examined quarterly.Then, seemingly out of nowhere,dozens of cabbage-like bud galls ap-peared one spring. While I couldnot locate shrubs nearby from whichthey may have spread to these newhosts, they certainly must have beenin the region. It took another yearfor me to rear the adults and havethem identified as a new midge spe-cies in the genus Contarinia.

TOP: Desert holly (Atriplex hymenelytra) iswidespread across the deserts into Utahand New Mexico. Its normal white color isoffset in spring when large, bright pink,cotton-candy-like galls develop from someof its buds. • BOTTOM: Creosote bush (Larreatridentata) is favored by many plant galls.

Several rather innocuous look-ing shrubs also support gall insects.The relatively unassuming burro-weed and related ragweeds (Ambro-sia spp.) support at least seven spe-cies of gall midges, while the vari-ous saltbushes and shadscales (Atri-plex spp.) support eight species.Other desert plants that also sup-


port gall insects include: bladder sage(Salazaria mexicana), indigo bush(Psorothamnus arborescens), golden-bush (Ericameria cuneata), cheese-bush (Hymenoclea salsola), desertbroom (Baccharis sarothroides),brittlebush (Encelia spp.), Mormontea (Ephedra spp.), horsebrush(Tetradymia spp.), and catclaw (Aca-cia greggii).

In 2009, a midge was reared froma bristly tubular gall on catclaw that,according to Dr. Raymond Gagné, aleading gall midge taxonomist, wasso astonishingly different from allother known species, it would likelyrequire the description of a new ge-nus and, perhaps, a new family. Howmany other great surprises are outthere?

But of all the interesting desertplants that have drawn my atten-tion, it is the desert oaks that havedominated much of my field timethese last several years. Muller’s oaks(Quercus cornelius-mulleri) and Pal-mer’s oaks (Q. palmeri) have becomea fount of new discoveries. Not onlydo they share species more commonto their northern relatives, but theyalso host several recently discoveredspecies.

While there is a small cluster ofMuller’s oaks in Joshua Tree Na-tional Monument, a much larger andcontiguous stand of these oaks oc-curs in the Northern Santa RosaMountains just southwest of PalmDesert. These trees support severalspecies of cynipid wasps includingthe beaked twig gall wasp (Dishol-caspis plumbella), and the saucer gallwasp (Andricus gigas), more com-monly associated with related whiteoaks of the Central Valley and sur-rounding foothills.

Muller’s oaks, however, haveyielded several new species not seenelsewhere. Some of these may proveto be the elusive and currently un-known alternate generations ofknown species. Specifically, cynipidwasps exhibit a rare phenomenonwhere a spring bisexual generationalternates with a summer-fall gen-

ABOVE: At least 16 species of gall midges(flies) are found only on the creosote bush,including Asphondylia auripila, whichinduces a gall about the size of a golf ball.This midge occurs throughout the rangeof creosote bush in the southwest. Thisgall was ground and smoked by the SereIndians of Arizona. LEFT: The gall Asphon-dylia digitata, often described as lookinglike moose antlers, is commonly found oncreosote bush. • BELOW LEFT AND RIGHT: Thetiny creosote stem gall midge (about .15inch), and an equally small wasp (about.18 inch) emerging from a beaked twiggall. Both insect photographs by Peter J.Bryant.


eration of females only. Many spe-cies are currently known only bythe adults from one generation,while others may be completely new

TOP LEFT: Rhopalomyia sp. on rabbitbrush. This unique flower gall, a new species of Rhopalomyia, was discovered on two different speciesof rabbitbrush, one in the northern Mojave and the other in Northern California. • ABOVE LEFT: One of the strangest interrelationships in theMojave has one insect inducing a gall in the gall produced by another insect. Pictured here is Aciurina trixa on rabbitbrush with the endogallof Rhopalomyia bigeloviae protruding. • ABOVE RIGHT: Cooper’s boxthorn, Lycium cooperi, is widespread throughout the desert’s sandy androcky flats and washes east into Utah and Arizona. This shrub also supports a new species of stem-galling moth, Gnorimoschema sp.

to science. Such is the mystery thatsurrounds what I call the melon gall,the lobe gall, and the lemon gallfound only on Muller’s oaks thus

far, all of which I have reared foridentification.

Even though Palmer’s oaks areclosely 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 headof cabbage, was found in the eastern Mojave. • MIDDLE: Rhopalomyia sp. on desert broom (Baccharis sarothroides). The galls of this newspecies of Rhopalomyia are heavily parasitized (94%) by torymid wasps in one area in the northeastern Mojave. • RIGHT: New gall specieson catclaw (Acacia greggii). The life cycle of this midge and many other gall insects in the desert is controlled by major rain events thatinfluence the development of new buds.


(Quercus chrysolepis) and huckle-berry oak (Q. vaccinifolia) and sharesome of their gall associates, Palmer’soaks support an interesting groupof largely unknown new species,unique to them. While five specieshave been reported on Palmer’s oaks,five additional new species were dis-covered just in 2009.

What drives my seemingly end-less quest for new plant galls is the

TOP: The beaked twig wasp gall (Disholcaspis plumbella) occurs on scrub oak (Quercusberberidifolia) elsewhere in the state, but also occurs on Muller’s oak (Q. cornelius-mulleri)in the desert. • MIDDLE LEFT: New species of bud gall on Muller’s oak. The galls of this newspecies produce males and females in the late spring, which suggests there is an alternategeneration later in the summer consisting only of females. • MIDDLE RIGHT: The lobe gall,a new species, was first found in the northern Santa Rosa Mountains above Palm Deserton Muller’s oak in 2009. Normal anthocyanin pigments found in plant tissue may beconcentrated in specific areas, as in certain galls, where the red-orange is expressed. Howand why this happens and what role the gall insect plays remain a mystery. Photographby Joyce Gross. • BOTTOM: Plant gall seekers are continually rewarded with new finds, suchas this small, green, “lumpy” gall discovered by the author in 2010.

perhaps unanswerable question ofhow many other species might bediscovered with a concentrated,long-term effort. The notion thatPalmer’s oaks, Muller’s oaks, anddesert scrub oak (Q. turbinella) aswell as other desert shrubs and treescould support dozens of new spe-cies, as well as unknown alternategenerations, is the magnet that con-tinues to lure me back.

My work in the desert over allthese years documenting the occur-rence of gall-inducing insects andmites has not been what I would callearth-shattering, but it does reveal atiny facet of desert life not previ-ously well understood. When youconsider a shrub like rabbitbrush orcreosote bush and take into accountall of the associate species depen-dent on it—the pollinators, leaf-eat-ing caterpillars, leaf miners, nectarthieves, and the complex mix of sev-eral species of gall insects—you havea biological universe within a singleshrub.

Then when one considers theramifications of the presence of asingle species of gall insect with allof its predators, parasites, their para-sites, and the insects that eat galltissue exclusively, the importanceof these species in the grand web ofdesert life staggers the imagination.By recognizing this tiniest of threads,we might just better understand thewhole fabric of desert life. Clearly,the biological diversity and the na-ture of interrelationships in thedesert are far more complex than wecurrently understand. And it is thiscomplexity that makes the risk oflosing species and systems that wedon’t even know exist that muchmore critical and irreversible.

A couple of years ago I foundmyself standing next to a Palmer’soak, at the head of a canyon in lateMarch, buffeted by a stiff, cold wind,chilled and hungry, wondering whatI was doing there. And then a small,green, lumpy gall I had never seenbefore caught my eye. After all theseyears, the Mojave and Sonoran


THE PALMER’S OAK MYSTERY

ne of the grand mysteries that has puzzled me for years, untiljust recently, is the occurrence of the same cynipid gall wasps

on Palmer’s oaks that are widely separated by hundreds of miles. Iexamined three Palmer’s oaks, for example, at the head of ShortCanyon in the Owen’s Peak Wilderness on the eastern side of thesouthern Sierra Nevada Mountains that were reported to me back in2008 by Joyce Gross (who works for the U.C. Berkeley NaturalHistory Museums and is an avid wildlife photographer).

These three trees stood by themselves, with the nearest Palmer’soaks over 100 miles away to the south in the southern New YorkMountains. The next nearest stand of these oaks occurs in the SantaRosa Mountains, southwest of Palm Desert. Yet, all three stands ofoaks share exactly the same species of cynipid wasps. How could thishappen? There is no way that these weak-flying wasps could havemigrated across the vastness of the Sonoran and Mojave Deserts andcolonized these new hosts, so how did they establish themselves ontheir hosts?

The most likely answer came with the discovery of a clone ofPalmer’s oaks in the Jurupa Hills, just east of Mira Loma and north ofthe 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 estimatedto have started growing during the late Pleistocene Era over 13,000years ago. These researchers think that climatic conditions at thetime were more favorable to the growth of Quercus palmeri and thattheir range was more contiguous, at least in Southern California.

With a warming trend developing since the last glacial period, therange of this oak and perhaps other relic populations like singleleafpinon (Pinus monophylla) have dramatically shrunk, leaving small,disjunct groups of these trees scattered in California. The idea ofPalmer’s oaks existing in a more contiguous range is supported by theoccurrence of the same species of cynipid wasps now found in theserelic groups of trees. Had Palmer’s oaks existed in a broader range,such as from the interior mountains of Southern California acrosswhat is now the Mojave Desert to the southeastern Sierra NevadaMountains, then the occurrence of the same species of cynipid waspsfrom west to east seems more plausible. Such is the case with theoccurrence of the exact same species on blue oak (Q. douglassii) alongthe Central Valley foothills from Redding to Bakersfield.

The small, dependent populations of gall wasp species on Palmer’soaks have existed generation after generation for thousands of yearson 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 travelinto space, the grand vastness of the unknown remains nearby in theMojave and Sonoran deserts.

Deserts remain the “New World” tome, revealing exciting discoverieswith each trip.

The discovery of new plant gallsin the Mojave is a reminder of how

Five new gall species have been discoveredon Palmer’s oak since 2009. Palmer’s oakfuzzy gall wasp (Trichoteras burnetti) andLyon’s gall wasp (Heteroecus lyoni) (TOP AND

MIDDLE) were previously known, but thehorn gall (BOTTOM) is a new species.

much we still have to learn aboutthe fabric of desert life.

Ron Russo, 4960 E. 12th Drive, Belling-ham, WA 98226, [email protected]

O


LEARNING TO READ A LANDSCAPEby Phil Van Soelen

both in the wild and in one’s owngarden. As a naturalist and gardener,that has become second nature tome, but it is a skill anyone can learn.It also requires some understandingof natural history—of phenomenasuch as sun exposure, landforms,and drainage patterns. Sometimesat the nursery I talk with peoplewho have little idea what the sundoes in their garden, or how thatchanges through the seasons. It’salso important to know the con-tours of your land and how it drains.You probably won’t make wise gar-dening decisions without thatknowledge.

As a society we have generallylost this skill and knowledge, largelybecause so many people lack anysort of personal relationship withnature. Acquiring these skills wasnecessary for our ancestors; their

LEFT: Ferns, manzanitas, dudleyas, and the winter’s flowing water in author’s drainageditch. These native plants are chosen to function as and to evoke a wild seasonal streamin 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 (Dudleyabrittonii), red fuchsia flowering gooseberry (Ribes speciosum), yellow western buttercups(Ranunculus occidentalis), and baby-blue-eyes (Nemophila menzesii) bloom on this seasonallymoist bank in the author’s garden. All exist with infrequent to no summer irrigation.Photograph by Saxon Holt.

survival often depended upon them.For instance, by observing, our an-cestors were able to recognize drain-ages where water persists just belowthe surface in the dry season. Thiswould have allowed them to spotthe combination of soil, moisture,and exposure to the sun that favorsedible roots and bulbs or that mightprovide a life-giving drink of waterin times of need.

So how does one develop suchskills in today’s world? Over time Ihave taught myself to hone my ownobservational skills for native plantgardening by “reading” the land-scape in the wild. I attempt to un-derstand the patterns I observe innature and to cultivate the patienceto look deeply by questioning whatI see.

For instance, why does a par-ticular species grow on a ridge as

Being able to “read” a landscapeis my personal goal whengardening with natives. Inpractice, this means placing

plants in locations that approximatetheir native and natural needs. Do-ing so greatly increases their chancesof survival over time—and eventhriving—without the constant in-put of water or other resources suchas fertilizers and pesticides. If a plantwe’ve placed in our garden thriveswithout too much attention fromus, then we’ve done our homeworkwell.

WHAT IT MEANS TOPONDER

To read a landscape successfullyrequires the ability to patiently ob-serve what is happening over time,


opposed to lower down? Sooften in California you willsee a hillside where there isoak woodland, then grass-land, and then chaparral.Through years of experienceobserving wild landscapes, Iknow that the oak woodlandis growing where there is adeeper soil profile with lesssun exposure, whereas thechaparral is usually growingin more rocky soil and hasa high sun exposure. Andgrasslands are somewhere inbetween. One of the things Ienjoy about hiking is con-stantly watching the land-scape change as you walkthrough it, and trying to in-terpret what is going on.

This ability to ponderwhat is happening in naturewas a skill that was honedthroughout human history,but today only a compara-tively few develop it. In na-ture, I find myself encour-aged to ask questions aboutwhat causes whatever it isI’m seeing. Why does a plantgrow there? What do the bees orhummingbirds feed upon in thespring versus in the summer or fall?Are birds making use of my garden?How do the roots from differentplants compete with each other?How does cold fall down on theland from above? How does waterflow across the land? Our gardensencompass an incredibly complexsystem of such interactions.

Ultimately, reading a landscapeis about developing a relationshipwith nature, a process I find satisfy-ing and that eventually becomes sec-ond nature. Once one begins to ac-quire this observational knowledge,it proves extremely helpful in guid-ing one’s gardening choices aboutwhat plants to select and where toplace them in the home garden. Thesame process can then be used toevaluate how well they respond, andto modify future gardening choices.

APPLYING THE SKILLS

For example, in my Sebastopolgarden, the previous owners had in-stalled French drains that channeledthe water into one corner of thegarden. This produced an ugly drain-age ditch, sometimes full of rushingwater in the winter, but bone dryall summer long. I had to contem-plate that as a design challenge. Howwould I use plants to soften the ditch,to camouflage it?

My solution was to create a fern-lined seasonal stream by plantingCalifornia polypody (Polypodiumcalifornicum)—a native fern one seesin nature along streams and canyonwalls, and that goes summer dor-mant—on both sides of the drain-age ditch. When the ditch dries upeach summer, the fern tolerates thatperfectly. I also planted several man-zanitas (Arctostaphylos manzanita x

Arctostaphylos densiflora)—natural hybrids of the com-mon manzanita and theVine Hill manzanita—onthe edge of the ditch, a spe-cies that is tolerant of mois-ture, good drainage, andsummer dryness. What I’veended up with is a lovelyseasonal creek of manzani-tas and ferns that is welladapted to the changes ofthe seasons.

Another plant I’ve uti-lized in my garden is adudleya hybrid (Dudleya x‘Frank Reinelt’, also knownas ‘Anacapa’) from theChannel Islands. It handlessome of same issues in adifferent way. As a succu-lent, it tolerates our sum-mer drought fine, butplumps up during the win-ter rains. I now have largedrifts of this in much ofmy garden. It has created alovely coastal bluff effectthat needs virtually no sup-plemental irrigation.

OFFERING CUSTOMERSADVICE

At the native plant nursery I co-own in northern California, custom-ers often ask us for advice regardingwhat is the right plant for a particu-lar site. Over time we have devel-oped a series of questions we askthem in order to guide their choices(see sidebar). These are the samequestions I almost unconsciously askmyself when placing plants in myown garden. For instance, custom-ers may tell us their garden is prima-rily in shade. Most plant labels onlydifferentiate between shade, partialshade or sun, or full sun. At firstglance it would seem a fairly easydecision to place shade-loving plantsin a shaded part of the garden. But totruly read a garden landscape in or-der to know what to plant and where

Native animals are often seen as pests, but they can functionin a native plant garden much as they do in the wild, with acertain beauty and harmony.


to plant it, we need considerablymore information than that.

There are different degrees ofshade, such as bright, high-canopydeciduous shade, versus deep ever-green forest shade. We need to knowwhat type of shade exists in thatparticular spot in the garden. Thereare a number of other factors thatalso should be taken into accountbefore making any planting deci-sions, such as the amount of avail-able moisture, the type of soil andavailable nutrients, the proximity tolarge trees or shrubs, how muchwind the site gets, and the angle andamount of sun exposure in differentseasons.

So, for example, if a customersays she would like to plant trilli-ums or lilies in her garden—plantsthat want moisture and brightshade—we need to locate the low-

SOME QUESTIONS TO GUIDE THE NATIVE PLANTGARDENER

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 blockingthe sun? Does it have open or dappled sunlight? What are itsqualities of shade—bright, deep, etc.—and what is creating theshade?)

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 majorpest 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 ofwhether it contains primarily clay, sand, or a mixture of the two, soyou can help gardeners make intelligent plant choices based ondrainage and soil types.)

6. Do you want primarily natives, or natives and non-natives that aredrought tolerant?

7. Are you comfortable with silver-gray foliage, or do you prefergreen and dark-green leaves? (This will have a major impact interms of the plants I recommend.)

8. Do you like things very tidy and formal, or more naturalistic andinformal?

Many of the same forces—substrate,nutrient flows, and exposure to sun—thatoperate on a larger scale in the homegarden, also stratify and segregate vege-tation patterns on the micro level, in thiscase lichens on a rock at the PinnaclesNational Monument.

est garden spot where water col-lects naturally, and determine if italso provides other requirementsof these plants, namely bright shadeyet little or no root competitionfrom other large plants or trees.Knowing what a plant requires foroptimum health and being able tofind the optimal location for it inthe garden are key to creating a


beautiful and successful native plantgarden.

Observing one’s own gardenkeenly throughout the seasons alsoreveals valuable information. As anexample, many shade tolerant plantsare deciduous (they lose their leavesseasonally) or herbaceous (they godormant back to their roots). Theysurvive the darkest days of winter in

a dormant state and leaf out just asthe angle of the sun raises enough toprovide adequate light for photosyn-thesis. This allows planting in areasdarkened by the low angle of the sunin the winter—more so than manyevergreen plants would tolerate—yetit still allows them to receive brightshade or sun in summer.

There are many reasons people

Sun exposure, drainage patterns, existing vegetation, and soils all combine to make a rich and pleasing mosaic on Mount Diablo. Natureis the best instructor in planning our home gardens.

garden. But for me, perhaps the mostcompelling is being able to recreateoutside my home the appealing inti-macy I sometimes experience whenI’m in the wild, the feeling that “thisspot feels so right.” The ability toread a landscape makes this possible.

Phil Van Soelen, 7560 Brittain Avenue,Sebastopol, CA 95472, [email protected]


DEVELOPING A SUSTAINABLE MIX FOR SEEDGERMINATION USING LOCAL MATERIALS

by Jackie Bergquist Salas, Michelle Laskowski, Brianna Schaefer, and Betty Young

Peat moss has been a major compo-nent of both media. Unfortunately,peat moss—one of the most com-mon amendments in germinationand growing media—is harvested ona scale much faster than peat bogsreplenish themselves and as such, isnot very renewable (Priesnitz 2007).Additionally, peat moss is harvestedand shipped from Canada. Coconutfiber (coir), another substitute forpeat moss, is shipped thousands ofmiles to reach our nurseries in theSan Francisco Bay Area. Shippingeither peat or coir uses a largeamount of fossil fuel.

Due to these ecological and en-vironmental concerns, the regularpotting medium used at the Nurser-ies was converted to a compost-based

blend produced within the park in2003. With careful control of thecompost quality, transplanted andmature native plants have performedvery well in this compost-based mix.However, so far we have been un-successful at formulating a seedlinggermination medium.

For the past few years, with fi-nancial support from the PresidioTrust, we have been trying to de-velop an alternative seed germina-tion medium with a rate of germina-tion similar to that of the commer-cial peat moss-based medium, whileusing local sustainable amendments.

In 2009, after a review of experi-mental literature, we chose compost,partially boiled rice hulls (a by-prod-uct of California’s agriculture indus-

TABLE 1. GERMINATION TRIAL MEDIA RECIPES

Trial Medium #1 Trial Medium #2 Trial Medium #3Ingredients

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), includesCanadian sphagnum peat moss, fine perlite, gypsum, dolomitic lime, and a low fertilizer charge. Results of ourtrial media were measured compared to this mix.

he Golden Gate NationalParks Conservancy’s NativePlant Nurseries are chargedeach year with the produc-

tion of 150,000-200,000 nativeplants grown from watershed-spe-cific seed, which are used in restora-tion projects throughout this na-tional park. The Nurseries’ goal is toproduce plants in the most ecologi-cally sustainable manner possible.This commitment includes consid-ering the ecological footprint of eachproduct used in plant production.

The main products used duringthe production process are seed ger-mination media, used in 16" x 16" x2" flats, and potting media for grow-ing seedlings that have been trans-planted into their final containers.

T


try), and earthworm castings aslocal and sustainable materials thatcould potentially provide the basicproperties of a successful growingmedium (Hidalgo et al. 2009,Hoitink et al. 1997, Evans R.Y. 2006,Merhaut 2006, Evans M. 2008,Merca 2009). Since performance ofrice hulls in germination media wasunknown, perlite was added as aporous/structural component tomaintain sufficient aeration in themix.

Media mixtures were preparedto test the amendments in differentcombinations (see Table 1, Germina-tion Trial Media Recipes). Samplesof the media were then sent to anindependent laboratory for analysisof their physical and chemical prop-erties. Flats were sown with threespecies of seed chosen for their vary-ing characteristics: California aster(Symphyotrichum chilensis, alsoknown as Aster chilensis), blue wildrye (Elymus glaucus), and stickymonkeyflower (Mimulus aurantia-cus). These seeds were sown in ran-dom on standardized and grid pat-terned germination flats that re-ceived intermittent mist set at a tem-perature of 65�F (18.33�C). Flatswere monitored to note the date ofgermination, potential mortality,and/or the date seedlings were largeenough to transplant. This data wascollected three times a week forseven weeks.

The results of the 2009 trial showthat the control peat-based mixtureproduced the highest germinationrate, fewest weeds (germinating fromweed seeds in the various amend-ments), lowest overall mortalityrates, and largest fresh and dryweight of roots and shoots. TrialMix #1, which was equal parts earth-worm castings, compost, and per-lite, and Mix #3 comprised of theseingredients plus boiled rice hullsseemed to perform nearly equallywell. Mix #2, equal parts rice hulls,compost, and perlite, lacked nutri-ent and water holding capacity, anddid not perform well in comparison

TOP: Developing a sustainable mix for seed germination using local materials. Left to right:Mix #1 (earthworm castings, perlite and compost), Mix #3 (earthworm castings, perlite,compost, and boiled rice hulls), Mix #2 (compost, boiled rice hulls and perlite). Allphotographs by Jackie Bergquist Salas. • MIDDLE: Flats of each mix planted with threenative species, including sticky monkeyflower, blue wild rye, and California aster, sixweeks after sowing. Clockwise from top left: Mix #3, Mix #2, Mix #1, and Control. •BOTTOM: Preferred experimental mix: Mix #1 after seven weeks.

to the other media evaluated(see Table 2, Results). Al-though the control mediumperformed best in terms ofpercentages, statistical data re-vealed no significant differ-ence between the germinationrates of Mix #1, #3, and thecontrol medium. However, thegermination rates in Mix #1and Mix #3 were only around75% of the germination rateof the control peat-based mix-ture.

In regard to physical prop-erties, Mix #1 was dense andbecame water logged, whileMix #3 with compost and ricehulls had better drainage, butdid not supply nutrientsquickly enough to the seed-lings, delaying growth. Theearthworm castings in Mix #1and #3 contained weed seedsthat had to be pulled. Whilethe rice hulls break downslowly, holding their shape in


the medium, the absence of earth-worm castings in Mix #2 created anutrient poor mixture for develop-ing seedlings.

The results of this trial show thatwith minor adjustments, compost,earthworm castings, and partiallyboiled rice hulls can perform wellwhen combined to make a germina-tion medium. Our next experimentwill be to modify Mix #3 with onepart compost, one part earthwormcastings, and two parts boiled ricehulls. This increase in rice hullsshould improve drainage, and theexclusion of perlite will make it moresustainable. We will sift the amend-ments through 1/4" hardware clothto provide a more uniform mix andadjust the fertilizers slightly for aquicker release.

We demonstrated that it is pos-sible to make local sustainable re-sources compare favorably with con-ventional non-sustainable products.With some simple modification tothe recipes, we are confident thatwe will soon be able to produce asustainable germination medium forour Nurseries. We would appreciate

TABLE 2. RESULTS OF EXPERIMENT BY SPECIES AND MIX

GerminationMortality

NumberRoot Root Shoot Shoot Root:

Averages%

% ofHarvested

Fresh Dry Fresh Dry Shootseedlings 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

hearing from other nurseries orgrowers about how they have madetheir media more sustainable.

REFERENCES

Evans, R.Y. 2006. Compost as a pot-ting mix amendment: Too much of agood thing? California OrnamentalResearch Federation News 10(3): 12.

Evans, M. January 2008. Rice hulls 101,Grower Talks. Ball Publishing, Chi-cago, IL.

Hidalgo P., M. Sindoni, F. Matta, andD.H. Nagel. 1997. Earthworm cast-ings increase germination rate andseedling development of cucumber.Mississippi Agricultural and ForestryExperiment Station, MSU Cares Re-search Report 22, No. 6. AccessedJune 18, 2009. http://msucares.com/pubs/researchreports/rr22-6.htm.

Hiotink H.A.J., M.A. Rose, and R.A.Zondag. 1997. Properties of materi-als available for formulation of high-quality container media. Ohio StateUniversity, Ohioline Bulletin, Orna-mental Plants Annual Reports and Re-search Reviews Special Circular 154.Accessed June 18, 2009. http://ohioline.osu.edu/sc154/sc154_14.html.

Merca, D. 2009. Personal communica-tion. California Rice Commission,Sacramento, CA.

Merhaut, D.J. 2006. Substrates and theiruse. California Ornamental ResearchFederation News 10(3): 1-2, 11.

Nelson, D.V. 2003. Chapter 6 in Green-house Operation and Management.Pearson Education, Inc., UpperSaddle River, NJ.

Priesnitz, W. 2007. Does peat mosshave a place in the ecological gar-den? Natural Living Magazine. Ac-cessed January 26, 2010. http://www.naturallifemagazine.com/0712/asknlpeat.html.

Jackie Bergquist Salas, Children’s Fairy-land, 699 Bellevue Avenue, Oakland, CA94610, [email protected];Michele Laskowski, Golden Gate NationalParks Conservancy, 1216 Ralston Avenue,Presidio of SF, San Francisco, CA 94123,[email protected];Brianna Schaefer, Golden Gate NationalParks Conservancy, 1216 Ralston Avenue,Presidio of SF, San Francisco, CA 94123,[email protected]; BettyYoung, Golden Gate National Parks Con-servancy, 201 Fort Mason, San Francisco,CA 94123, [email protected]


NEW CNPS FELLOW: ROGER RAICHEby Phyllis Faber

oger Raiche has made unusu-ally extensive contributionsto California botany in fourdistinct arenas. As a foremost

California botanist, he has gener-ously shared his extensive knowl-edge with many audiences, includ-ing numerous members of CNPS,and has discovered several new na-tive species that have subsequentlybeen named for him. As a gardenerpar excellence, for years he main-tained the California Native sectionof the University of California Bo-tanical Garden at Berkeley, and de-

veloped an extraordinary serpentinegarden.

As a horticulturalist, Roger madeextensive collections of native seed,always with precise collection data,collected numerous plant variants,and developed several interestingcultivars for gardens. And as a landprotector of the best sort, he pur-chased a core area of “The Cedars,”an extraordinary serpentine barrenin Sonoma County in order to pro-tect it. He has been host to manyCNPS members and to fascinatingand delightful chapter field trips to

The Cedars. In sum, Roger’s life hasbeen devoted to California’s floraand to sharing his love for this flora.

Roger worked for 23 years at theU.C. Botanical Garden (UCBG) inBerkeley in charge of the 15-acreCalifornia Collection. He was theprimary collector of that section forall those years and brought in thou-sands of wild plants during that timewith precise collection data. He wasalso primary collector for the Gar-dens’ Index Seminum or seed listthat is put out every two years andgoes to over 500 botanical institu-

Roger in 2011 at The Cedars with his beloved dogs, Serpentino (gray), Hortus (black), and Daisy (tan). The Cedars has been a focus ofRoger’s life since 1981 when he first entered its canyons. Photograph by David McCrory.

R


tions around the world (for a totalof ten lists during his tenure at theGarden). Although most of hiscollecting was done for UCBG, heconducted most of it on his owntime.

Roger wrote a few articles overthe years pertaining to natives, in-cluding an article in Pacific Horti-culture in 1991 describing many ofhis cultivars. He also wrote a col-umn called the “California Corner”for a monthly internal UCBG news-letter called the UC Bee, where hewrote about various things of inter-est happening in the California area.Roger was also on the Jepson ManualHorticultural Council, which helpedprepare the horticultural entries forThe Jepson Manual (1993) that War-ren Roberts chaired. Roger com-

mented on numerous plant entriesin that volume.

Perhaps Roger’s most impressivetalent regarding native plants washis ability to find them, especiallyrare ones in the wild, as well asfinding new ones. Three new taxawere named in his honor: Cedarsfairy-lantern (Calochortus raichei),Raiche’s red ribbons (Clarkia con-cinna ssp. raichei), and Raiche’s man-zanita (Arctostaphylos stanfordianassp. raichei). Roger was also verytalented at finding variants that hethought might make interesting cul-tivars for horticulture, includingaround 20 that he named. Roger’sintroductions of exceptional nativeplant cultivars frequently showed upfor sale at the CNPS East Bay Chap-ter’s plant sales before they had re-ceived wider circulation. This cer-tainly had the effect of supportingthe chapter by making the sales moreinteresting.

Several native genera have beenof special interest to Roger, espe-cially Arctostaphylos, Ceanothus, andStreptanthus. He made many hun-dreds of herbarium specimens ofthose and other taxa over the years.This work helped to identify manynew sites for existing and possiblysome new taxa. In recent years,Roger collaborated with Dr. JamesReveal of Cornell to publish twonew taxa that occur at The Cedars,The Cedars buckwheat (Eriogonumcedrorum) and The Cedars ocean-spray (Holodiscus dumosus var. cedro-rus).

Monocots also really interestedhim, and he was a major contributorto the CNPS book, Wild Lilies, Irises,and Grasses: Gardening with Califor-nia Monocots, finally published in2003 after 30 years of a workingstudy group. Roger did the primarywriting for Brodiaea, Dichelostemma,Trietleia, Fritillaria, and all theGrasses, Sedges and Rushes—a ma-jor component of the book.

One of Roger’s major obsessionshas been the serpentine flora of Cali-fornia, and much of his collecting

has focused on plants growing onthat substrate. Protecting and un-derstanding an area in SonomaCounty called The Cedars becamepart of his obsession, and he becamea vigorous advocate for its conser-vation since 1983. He has led manyfield trips for CNPS to this site, aswell as giving numerous slide lec-tures over the years pertaining tothe native flora growing on this hugeserpentine area which has beeneroded into many deep narrow can-yons. At least seven distinct plantsare found only at The Cedars.

In 1999, Roger and his partner,David McCrory, purchased 520 acresin the heart of The Cedars. Thisallowed them to restore damagedareas, to continue with field trips,and to facilitate scientific studies. In2011, in a collaborative effort withSave The Redwoods League, the StateCoastal Conservancy, and the Gor-don and Betty Moore Foundation,500 acres of this important core areawere purchased and transferred toBLM (Bureau of Land Management)to add to its Area of Critical Envi-ronmental Concern at The Cedars.Roger also helped the Sonoma LandTrust (SLT) develop its “Conserva-tion Plan for The Cedars Area,”which will guide future conserva-tion efforts of this region.

Warren Roberts, Superintendentof the U.C. Davis Arboretum (nowretired), and former president of theCalifornia Horticultural Society, de-scribed Roger as follows:

Roger Raiche fell in love with theplants of California literally at firstsight. Driving across the Sierra enroute from his home town, New-port, Rhode Island, to his new lifein California, he was struck by thebeauty of our native manzanitas.He never got over it. His enthusi-asm for the flora of his adoptedhome state has grown over theyears, and for decades he has beenone of California’s greatest advo-cates for the beauty and garden-worthiness of our native plants.

Roger with his business partner, DavidMcCrory, at The Cedars. Photograph byPhil Van Soelen.


Phil Van Soelen, Cal Flora Nurs-ery co-owner in Santa Rosa, and apast president of the Milo BakerChapter of CNPS, made the follow-ing observation of Roger:

Roger really is a towering figure inhorticulture in general, and Cali-fornia native plant horticulture inparticular. I believe he took themantle directly from WayneRoderick. A state CNPS surveyshowed that, among all respondentswho answered it, members’ inter-est in’gardening placed at the topof the list of topics. Certainly gar-dening with native plants wasfirst developed and showcased tothe public in botanic gardens, andRoger’s contributions in that fieldare great. There really is no oneelse of his caliber and accomplish-ments. While he was at the Berke-

ley Botanic Garden, the Californianative area was transformed from anice, rather unexceptional gardenof natives to a very dynamic artisticand scientific creation. Roger has avery rare blend of exceptional ar-tistic abilities as well as acute sci-entific perception and memory. Hehas always been extremely gener-ous with his time and energy lead-ing CNPS field trips. My businesspartner Sherrie Althouse said thatRoger led the best field trips shehas ever been on, and from myexperience I would agree. His tripsto the Cedars are legendary.

Roger Raiche has perhaps beenone of the finest and most effectivenative plant advocates of this gen-eration, and California has been for-tunate to have his skills, energy, anddevotion in understanding and ap-

preciating our native flora, making itknown to others, and protecting it.

Phyllis M. Faber, 212 Del Casa Drive,Mill Valley, CA 94941, [email protected]

A vintage photograph of Roger back in 1987, standing in a chaparral (on Cow Mt. east ofUkiah) of the Raiche manzanita (Arctostaphylos stanfordiana ssp. raichei). The species wasnamed in his honor that year by Walter Knight of the California Academy of Sciences.Photograph by Roger Raiche.

Two of the 20 cultivars that Rogerhas propagated and named. TOP:‘Joyce Rose’, a deep rose selectionof the coast flowering currant,Ribes sanguineum var. glutino-sum, which he named in honorof his mother, Joyce Rose Raiche.MIDDLE: The magnificent burgun-dy leaf stream orchid, Epipactisgigantea f. rubrifolia ‘SerpentineNight’. Both photographs by Phil VanSoelen. • BOTTOM: Roger has demonstratedan uncanny ability to find rare and newnative plants in the wild. One of three newtaxa named in his honor is The Cedarsfairy-lantern (Calochortus raichei). Photo-graph by Roger Raiche.


SCOTT FLEMING: 1923–2011by John Danielsen

cott, with his wife Jenny, wasamong the founding mem-bers of CNPS in 1965, andbecame a Fellow of the Cali-

fornia Native Plant Society in 1985(see the April 1985 issue of Fre-montia). Scott was active in CNPSat the state level as treasurer andlong-time legal advisor, where he

brought creativethinking to solvethe Society’s earlyfinancial growingpains, as well asnegotiating thecontract for CNPSto produce its firstplant poster.

A lawyer bytraining, Scottused his skills toput together, withothers, the Kaiser-P e r m a n e n t eHealth Program,

and helped to found the Planningand Conservation League, where heworked tirelessly promoting conser-vation activities in California. Scottwas an avid white water kayaker whoalso enjoyed hiking and camping.Scott and Jenny are survived bydaughters India and Hilari as well asthree granddaughters.

I first met Scott on one of ourchapter’s weekend hikes, hoping todiscover the wonders of the BayArea’s native flora. We also discov-ered that we worked for the samecompany, Kaiser-Permanente ofOakland, and that our spouses weregood friends. Scott and Jenny were awonderful and inspiring team whowarmly welcomed my wife Charliand me into their home and family.

They continued over many de-cades to provide this hospitality toother CNPS members from all overthe state who were in need of a goodsleepover while attending the many

chapter and state CNPS board meet-ings, or conservation strategy andplanning sessions held in Berkeleyor hosted at their home.

Scott loved to travel on field tripsto the far corners of the state, andwe shared this passion with Scottand Jenny. Scott also enjoyed shar-ing meals and conversation withfriends, and he was an expert chefwith his famous in-house smokeoven. When Scott decided to stepdown as state CNPS treasurer, heasked me to take on the treasurerposition. He was always ready toprovide good and insightful help asCNPS nearly doubled in size overthe next few years. He was equippedwith that unique quality to be ableto listen carefully to what the prob-lem of the moment appeared to be,rephrase it in a way that helped oth-ers understand, and then suggestcreative solutions.

I often think of a raft trip wetook down the Colorado River inthe early 1990s. Scott and I hostedabout 25 people on a two-week rub-ber raft trip through most of thewhite water available to boaters. Wetraveled in small rafts, each holdingabout five persons. At one point onthe river we were faced with mon-strous waves going over Lava Falls.Observing these from the banks ofthe river, one person commented,“Holy smokes, I am not going overthose,” to which Scott replied,“Watch me, I’m going over them ina kayak!” Well, we all watched Scottmake it, so the rest of us followed,inspired by his example. Scott wascapable, fearless, and expert on thesetrips, which were wonderful quali-ties to have in a friend and colleague.

Scott and Jenny loved to garden,although truth be told, Scott spentsome of the garden time doing kayakrollovers in the family pool. WhileScott was the prime builder of the

garden infrastructure, Jenny took thelead in acquiring the plants. Throughthis teamwork they created a homegarden that remains one of the sig-nature native plant gardens in theBay Area.

The effort to create their garden,plant by plant and rock by rock overmany years, also reflected the dedi-cated plant conservation ethic theyboth shared and engendered withinCNPS. And as we remember ourdear friends, one legacy of their lovelives on in the beautiful garden theygenerously shared with all.

John Danielsen, 10 Kerr Avenue, Kensing-ton, CA 94707, [email protected]

S

Scott and Jenny in their Bay Area nativeplant garden.

Scott Fleming.


RALPH MILTON INGOLS: 1911–2011by Leah Price Hawks

e have lost one of thefinest conservationiststhat one could everhave known. Centenar-

ian Ralph Milton Ingols passed awayin Napa, California on July 11, 2011.

A product of Weed Patch in KernCounty, California, Ralph was theyoungest of seven siblings. He andhis wife Evelyne were parents of the“Three J’s,” Janet, Jim, and JudyIngols, to whom they were devoted.Their children joined them in a life-time of outdoor activities, whichcemented their interest in the envi-ronment as well as in education.

As a toddler, Ralph was oftenput on a blanket among the wild-flowers and grasses in Weed Patch.His ground-eye view of the land-scape left a lasting impression onhim and was his earliest instructorin the natural world. At a young age,Ralph knew he wanted to becomean educator, and later on he did.

He received his B.A. and teach-ing credential at U.C. Berkeley andthen began a 35-year career—mostof which was spent at St. HelenaHigh School in the Napa Valley—teaching California and world his-tory, world literature, public speak-

ing, and journalism. During thistime he received his master’s degreeand counseling certification at SanFrancisco State University. Ralphcoached numerous sports includingfootball, basketball, baseball, wres-tling, and boxing. He also gave les-sons to the school ski club at hiscabin in the Sierra Nevada. Ralphand Evelyne were active in the NapaValley Chapter of the California Na-tive Plant Society, and for many yearsused their backyard to propagatenative perennials for the chapter’splant sales. After they both retiredin 1972, the couple spent many yearstraveling throughout California,studying and collecting its nativeflora and gaining valuable knowl-edge and background in plant iden-tification and propagation.

During this period, the Ingolsand their friends first began discuss-ing the creation of a public garden inthe Napa Valley where visitors couldlearn about and enjoy California’sflora in protected surroundings. Theidea of a garden that could be usedas an outdoor classroom particularlyappealed to the Ingols, since bothwere career educators.

By 1985, with the help of con-

servationist friends, the Napa ValleyChapter of CNPS, and the SkylineCitizens Association, they foundedthe three-acre Martha Walker Na-tive Habitat Garden, located at Sky-line Wilderness Park, and becameits first curators. Ralph and Evelynewere the driving force behind thenew garden in its beginning years,and Ralph continued as curatoremeritus after Evelyne’s death in2000.

Those close to Ralph knew thathe always had a broad vision for the

Ralph Ingols in the mid-1950s at St. HelenaHigh School, where he taught for over 30years.

W


In his later years, Ralph contin-ued to make every effort to try newthings at an age when most peopleconsider their life’s work finished.He wrote, planned, and organizedthe community to accomplish wor-thy goals, and in the process con-nected with others in every walk oflife.

Ralph always spoke warmly ofothers he respected, whether com-munity leaders, former students, vol-unteers, or youth, and never misseda chance to encourage others. Inone way or another, Ralph was amentor to everyone he met. He sim-ply felt that it was within the capac-ity of each person to succeed, andhe was capable of convincing gen-erations of students and friends thatthis was so.

Over the past ten years Ralphpublished two books for his family:The Ingols Cabin, stories of fam-ily and student adventures in theSierra, and Why I Am Who I Am, abiographical sketch of his life in Cal-ifornia. He was a prolific writer, and

also finished five other books dur-ing this time, two with coauthors,including one about St. Helena HighSchool, and another on Skyline Wil-derness Park. Proceeds from the saleof his books go to support CNPSand the Martha Walker Garden.

Just a few years ago, Ralph re-ceived the Earl Thollander Conser-vation Award, presented annuallyby the Napa Group of the SierraClub to an individual who hasachieved excellence in conservation.

Perhaps one of Ralph’s most im-pressive accomplishments was hishaving the Martha Walker Gardennamed a World Peace Garden in2002, the first in the United Statesto receive that honor! (Each gardenin the England-based World PeaceGarden Project serves as a retreatfor peaceful reflection on the beautyof nature and the ideals of living inharmony with others.) In 2010 Ralphattended the celebration of theGarden’s 25th anniversary, and upuntil a few years ago, he also at-tended as many class reunions of hishigh school students as he could.

Ralph was planning for the fu-ture up until the day before he passedaway, still making arrangements forthe Garden to benefit in yet moreways through plantings, work par-ties, and proceeds from his books.“When people go into the Garden,I want them to see butterflies every-where,” he said. “I want to plantDutchman’s pipevine for them.”

His friends in the Napa ValleyChapter are now working to makethat dream come true. Additionalmilkweed to attract monarch but-terflies and Dutchman’s pipevineto attract the pipevine butterfly arejust two of the species being dug innow, and a steering committee isplanning a Ralph Ingols ButterflyTrail to be completed in 2012 tohonor Ralph’s dedication and visionfor the Garden.

Leah Price Hawks, 1236 Second Avenue,Napa, CA 94558, [email protected]

ABOVE: The Ingols, both teachers, believedstrongly in using the Martha Walker Gar-den as an outdoor classroom. Ralph, oneof the garden’s science docents, had a knackfor sharing with students his love of plantsand animals. • LEFT: Ralph with his wife,Evelyne, in the Sierra Nevada in the mid-1990s. The two were the driving forcebehind the creation of the Martha WalkerNative Habitat Garden in Napa County.

Garden. “Future generations canenjoy what we prepare for them to-day!” was his motto. Thanks largelyto his efforts, the Napa Valley Chap-ter established a science docent pro-gram there. Today that program en-sures that young students who visitthe Garden learn about its plantsand wildlife.

Ralph spoke of a time, a hun-dred years from now, when thisgeneration’s grandchildren wouldalso walk the paths of the MarthaWalker Garden under his favoritetrees. He willingly gave whatever hecould in energy and resources to thecommunity, and in particular to theGarden. This was his legacy.


LETTERS TO THE EDITOR

WETLANDS: MISPLACEDEMPHASIS

Having studied the seasonal wet-lands of Sonoma County’s Santa RosaPlain (SRP) for the last 27 years (con-sulting to developers, city, county, andstate agencies, preserve proponents,nonprofits and landowners, and as acongressional appointee to the SonomaCounty Vernal Pool Task Force, 1988-1992), I am glad to see the attentionbeing paid to this unique area (seeFremontia Vol. 37, No. 4/Vol. 38, No.1, pp. 40-43). However, I feel com-pelled to comment.

First, with regard to history, itshould be noted that well before 1970,this region’s broad valley of seasonalwetlands, perennial grasslands, oaksavanna, and riparian forests hadalready been thoroughly (80-90%)chopped up, chopped down, con-verted to prunes, pears, grapes, dair-ies, mustard and annual grassland,and dewatered by roads, ditches,berms, buried drains, and engineeredchannels that flow straight to the La-guna de Santa Rosa. Up until the early1980s, federal law allowed for the“minor fill” of up to one full acre ofwetland under the “nationwide per-mit program,” often with little or nosignificant mitigation.

Then, with probably less than 5%of the region’s wetlands left, intenseefforts were made over the last severaldecades to reduce, modify, and/or miti-gate modern development’s wetlandimpacts. For 25 years now, the U.S.Army Corps of Engineers has requiredat least 1:1 replacement of wetlandfunctions, values, and acreage (essen-tially dictating new wetland construc-tion). In addition, the U.S. Fish andWildlife Service currently requires theprovision of acreage or credits sanc-tioned as “Establishment” habitat, i.e.,new physical habitat constructed andseeded with listed species.

The region’s initial efforts towardrare plant translocation (e.g., AltonLane) were conducted at the directionof, and in fact as a permit requirementof the California Department of Fishand Game. Hence, on the SRP, where

there is so precious little natural wet-land habitat left—probably well lessthan 5%, and even that is severely frag-mented—I believe we no longer evenhave the same underlying ecosystemwithin which to manage the remnants,and that continued habitat construc-tion/restoration with rare plant trans-location will likely be important partsof any hoped for recovery.

Wetland (and hence native poolflora) conservation on the SRP hasalready come a long way since the1980s, thanks largely to people likeBetty Guggolz and Nancy Harrison(both had been active members ofCNPS) and others, plus a more activerole by the agencies, specifically re-quiring compensatory mitigation. Theresults to date, while far from perfect,include the long-term preservation ofmore than 1,500 acres on more than50 preserves.

Finally, while I encourage all sci-entific efforts on the SRP, I also be-lieve that today’s main challenge—

as it has been for decades—is simplyto save as much suitable wetland-supporting land as possible (actualand potential), enhance it as needed,funded, and practical (including planttranslocation), and lock it away. Dowe need to improve our preserve man-agement? Absolutely. Do we need moreresearch? Always. I’m not sure if theseare desperate times calling for desper-ate measures just yet, but in some re-spects it feels very close.

Charlie PattersonPlant Ecologist, Lafayette, CA

NEED FOR SCIENCE-BASEDCONSERVATIONSTRATEGIES: THE AUTHORSREPLY

We appreciate Mr. CharliePatterson’s perspective in his responseto our article on the vernal pools of theSanta Rosa Plain in Sonoma County.


While the plants of the Santa RosaPlain are threatened by human im-pacts in many ways, they are alsodeeply valued by a large number ofpassionate local constituents that areeager to contribute to the conserva-tion of the flora. We are aware of Mr.Patterson’s experience with vernal poolmitigation efforts on the Plain, appre-ciate his genuine interest and concernfor vernal pool communities, and wel-come his outlook on our article. How-ever, we respectfully disagree that ouremphasis on science-based conserva-tion and community involvement onthe Santa Rosa Plain is misplaced, andwe are eager to reiterate our positionthat scientific research is an essentialpart of ensuring the long-term sur-vival of vernal pool habitat.

The additional history provided inMr. Patterson’s letter nicely comple-

ments our forward-looking perspec-tive on vernal pool conservation onthe Santa Rosa Plain. Indeed, habitatloss in the area began well before theearly 1970s, yet the rapid urban ex-pansion during 1970–1990 unequivo-cally pushed the threats facing nativehabitats in this region to a new level ofurgency. As discussed in our originalarticle, the accelerating loss of vernalpool habitat intersected with endan-gered species regulations during thistime to prompt the regulation of ver-nal pool mitigation projects by CDFGand other agencies. In the meantime, agreat deal of mitigation, translocation,and habitat loss occurred before thebasic ecological requirements of ver-nal pool species were understood.

We fully agree with Mr. Pattersonthat our main challenge is to save asmuch suitable wetland-supporting

land as possible. But we regard sci-ence as the most effective means todetermine what is “suitable.” Scien-tists estimate that only 5-10% ofCalifornia’s original vernal pool habi-tat remains, and a great deal of recentconservation efforts have been chan-neled into mitigation, while destruc-tion of natural habitat continues. Butif man-made wetlands fail to providesustainable conditions for translocatedpopulations, then we are faced withthe unfortunate reality that vernal poolmitigation may be a slow and expen-sive method for extirpating popula-tions, and eventually the species thatwe aim to protect.

Mr. Patterson, who has personallybeen involved in numerous mitigationefforts, admits that the results of re-cent conservation efforts may be “farfrom perfect.” But just how far? Arethey too far to be an effective conser-vation strategy? Hopefully not, but weargue that mitigation decisions—andtheir consequences—should be morecritically monitored, and more heavilyresearched. Vetted protocols, appro-priate controls, adequate replication,and rigorous data analysis can facili-tate the objective evaluation of the de-gree to which mitigated vernal poolsare supporting sustainable vernal poolpopulations and communities. Fur-thermore, detailed studies of remnantpopulations are essential for determin-ing the important biological processesinfluencing population growth andpersistence in the fragmented contextthat now characterizes the vast major-ity of historic vernal pool habitat.

The problem, of course, is thatthoughtful science and careful moni-toring takes time, and often must oper-ate at a slower pace than the pressuresof development and the rates of habitatloss. Fortunately, there are conserva-tion scientists willing to take that timein the Santa Rosa Plain. Hopefully, thepractitioners of vernal pool mitigationand conservation will facilitate that re-search by sharing any monitoring data,releasing historical records, and facili-tating site access, and will be eager todevelop strategies that incorporate theinsights from these projects.

Nancy C. Emery, Purdue University

Michelle Jensen, Ecologist,Pepperwood Preserve

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F R E M O N T I AV O L U M E 3 8 : 4 / 3 9 : 1 , O C T O B E R 2 0 1 0 / J A N U A R Y 2 0 1 1

❏ Enclosed is a check made payable to CNPS Membership gift:

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Join Today!

Please make your check payable to “CNPS” and send to: California Native Plant Society, 2707 K Street, Suite 1, Sacra-mento, CA 95816-5113. Phone: (916) 447-2677; Fax: (916) 447-2727; Web site: www.cnps.org.; Email: [email protected]

❏ Enclosed is a matching gift form provided by my employer

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CNPS member gifts allow us to promote and protect California’s native plants andtheir habitats. Gifts are tax-deductible minus the $12 of the total gift which goestoward publication of Fremontia.

NAME

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❏ $1,500 Mariposa Lily ❏ $600 Benefactor ❏ $300 Patron ❏ $100 Plant Lover

❏ $75 Family ❏ $75 International or Library ❏ $45 Individual ❏ $25 Limited Income

CONTRIBUTORS (continued from back cover)

CORPORATE /ORGANIZATIONAL

❏ $2,500 10+ Employees ❏ $1,000 7-10 Employees ❏ $500 4-6 Employees ❏ $150 1-3 Employees

Patrick Kobernus, biologist, supervised habitat management and endangeredspecies monitoring on San Bruno Mountain from 1995–2007. He is currently aconsultant for Coast Ridge Ecology, and a member of the Yerba Buena Chapter ofCNPS.

Michele Laskowski is seed ecologist for the Presidio Native Plant Nursery in theGolden Gate National Parks.

Ron Russo is the retired chief naturalist of the East Bay Regional Park Districtwhere he worked for 37 years, and the author of Field Guide to Plant Galls ofCalifornia and Other Western States.

Brianna Schaefer is manager of the Presidio Native Plant Nursery in the GoldenGate National Parks.

Hugh Safford is regional ecologist for the U.S. Forest Service’s Pacific SouthwestRegion, and a research associate with the Department of Environmental Scienceand Policy, University of California, Davis.

Phil Van Soelen is co-owner of California Flora Nursery, a past president of theMilo Baker Chapter (Sonoma County) of CNPS, and teaches classes at Santa RosaJunior College on gardening with California native plants.

Betty Young is director of the six native plant nurseries located throughout the80,000 acres of the Golden Gate National Parks in Marin, San Francisco, and SanMateo Counties.

MATERIALS FORPUBLICATION

Members and others are invitedto submit material for publica-tion in Fremontia. Instructionsfor contributors can be foundon the CNPS website, www.cnps.org, or can be requested fromFremontia Editor Bob Hass [email protected].

Fremontia Editorial AdvisoryBoard

Susan D’Alcamo, Ellen Dean,Phyllis M. Faber, Holly Forbes,Brett Hall, Tara Hansen, ToddKeeler-Wolf, David Keil, PamMuick, Bart O’Brien, RogerRaiche, John Sawyer, TeresaSholars, Greg Suba, Dick Turner,Mike Vasey, Carol Witham

F R E M O N T I A V O L U M E 3 8 : 4 / 3 9 : 1 , O C T O B E R 2 0 1 0 / J A N U A R Y 2 0 1 1

CONTRIBUTORS

California Native Plant Society2707 K Street, Suite 1Sacramento, CA 95816-5113

Address Service Requested

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U.S. PostagePAID

A.M.S.

FROM THE EDITOR

(continued on inside back cover)

Earl B. Alexander is a retired pedologist (soils) and activegeoecologist who has worked for several agencies, includ-ing the U.S. Forest Service in California, and investigatedserpentine soils and plant communities from Baja Californiato Alaska.

Jackie Bergquist is the landscape supervisor at Children’sFairyland in Oakland, CA.

Jim Bishop, with his botanist wife Catie Bishop, has con-ducted GLORIA field work since the project began in Cali-fornia in 2004. He also helped to develop the strategy andmethods for the project’s downslope surveys.

Daniel S. Cooper is an independent consultant based in LosAngeles. From 1999 to 2005 he worked for Audubon Cali-fornia. The author of Important Bird Areas of California(2004), he is a member of the LA/Santa Monica MountainsChapter of CNPS.

John Danielsen is a CNPS Fellow.

Stephen W. Edwards is the third director of the RegionalParks Botanic Garden in Berkeley, CA, and has served in thatposition since 1983.

Phyllis M. Faber is a past editor of Fremontia (1984–1999),and for many years also served as CNPS publications chair.

Leah Price Hawks was the editor for Ralph Ingols’ books,and is coauthor of Skyline Wilderness Park, Nature’s Gift toNapa Valley.

t is timely that Fremontia carry an article concerned withclimate change. Systematic observations of biological sys-tems are yielding important insights about how they are

responding to climate change. Measuring the ecologicaleffects of climate change requires field observations thatspan decades, not months or years. So it takes quite awhilebefore scientists are able to gather sufficient data in order todraw meaningful conclusions.

Jim Bishop’s lead article in this issue is about climatechange. In it, Jim introduces us to a fascinating interna-tional research effort—the Global Observation ResearchInitiative in Alpine Environments (known simply asGLORIA). As he explains, alpine environments are particu-larly conducive to measuring the biological effects of globalclimate changes. This is because human disturbance in thealpine zone is generally minimal, alpine environments spannearly all latitudes and elevations, and they include themajor climate zones of the world. The article then goes onto explain more about the alpine zone, and the environ-mental stresses on alpine plants and how they adapt.

The GLORIA project established its first field sites inEurope in 2001, and three years later the first field sites inthe Western Hemisphere were set up in California. Jim ispart of the California team of scientists and volunteers thatis taking surveys at monitoring sites using the standardizedinternational protocol so the results can be shared globally.

While data gathered so far is preliminary, it suggeststhat plant diversity is increasing in the alpine zone, whichwould be expected if average annual temperatures world-wide are warming. With GLORIA and similar projects, westand not only to learn about the response of alpine plantsto climate change, but to greatly expand our knowledge ofhigh-elevation vegetation in California and worldwide.

—Bob Hass

I

vol. 38, no. 4 and vol. 39, no. 1 • fremontia...volume 38:4/39:1, october 2010/january 2011...

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