VOL. 42, NO. 1 • JANUARY 2014 FREMONTIA...VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 1 CONTENTS THE COVER: Sunrise on the east side of the Granite Mountains in the Mojave National Preserve

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JOURNAL OF THE CALIFORNIA NATIVE PLANT SOCIETY

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VOL. 42, NO. 1 • JANUARY 2014

FREMONTIA

CALIFORNIA’S DESERTS, PART 1: BIOLOGY AND ECOLOGYCALIFORNIA’S DESERTS, PART 1: BIOLOGY AND ECOLOGY

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Staff and board listings are as of January 2014.Printed by Lithtex NW: www.lithtexnw.com

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.

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VOL. 42, NO. 1, JANUARY 2014

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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—oftheir ideas, statements, or opinions.

CALIFORNIA NATIVEPLANT SOCIETY

Dedicated to the Preservation ofthe California Native Flora

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

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

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

Lisa GiambastianiMojave Desert: Tim ThomasMonterey Bay: Brian LeNeveMount Lassen: Catie BishopNapa Valley: Gerald TombocNorth Coast: Larry LevineNorth San Joaquin: Alan MillerOrange County: Nancy HeulerRedbud (Grass Valley/Auburn):

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StephensSanta Clara Valley: Judy FenertySanta Cruz County: Deanna GiulianoSequoia (Fresno): Paul MitchellShasta: Ken KilbornSierra Foothills (Tuolome/Calaveras/

Mariposa): Robert BrownSouth Coast (Palos Verdes):

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Mary Frances Kelly-PohYerba Buena (San Francisco):

Ellen Edelson

Laura Camp (President); David Bigham(Vice President); Carolyn Longstreth(Secretary); Nancy Morin (Treasurer);At-Large: Kristie Haydu, Bill Hunt,Gordon Leppig, David Varner, MichaelVasey, Steve Windhager; ChapterCouncil Representatives: Orchid Black,Glen Holstein

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CONTENTS

THE COVER: Sunrise on the east side of the Granite Mountains in the Mojave National Preserve. In the foreground is a skeletonof buckthorn cholla (Cylindropuntia acanthocarpa) and cinchweed (Pectis papposa var. papposa). September 2013. Photographcourtesy of Stephen Ingram, www.ingramphoto.com.

EDITORIAL by Kara Moore ...................................................................................... 2

FLORISTIC DIVERSITY AND DISCOVERY IN THE CALIFORNIA DESERTby James M. André ......................................................................................... 3The California desert flora is surprisingly diverse and a hotbed for taxonomicdiscovery. Looming impacts, however, pose a major threat to this unparalleledfloristic frontier.

RARE PLANT DIVERSITY IN THE CALIFORNIA DESERTS: PRIORITIES FORRESEARCH AND CONSERVATION Kara A. Moore and James M. André ......... 9Many of California’s rare and unique desert plants are at risk, and research on theirbiology and distribution is urgently needed to fuel effective conservation strategies inour rapidly changing deserts.

APPLYING SPECIES DISTRIBUTION MODELING TOIDENTIFY RARE SPECIES HOT SPOTSby Patrick McIntyre ............................................................ 15

How researchers are using predictive mapping tools to understand plant diversity inCalifornia’s deserts.

THE CASTLE MOUNTAINS: A HOTSPOT OF DESERTPLANT DIVERSITY by Duncan Bell .................................. 17Recent explorations of the Castle Mountains in the Eastern Mojave show that they arerich in botanical diversity.

MICROBIOTIC SOIL CRUST COMMUNITIES: A CRITICAL COMPONENTOF CALIFORNIA’S DESERTS by Nicole Pietrasiak and Jeffrey R. Johansen ...... 18Microbiotic soil crusts are in dire need of conservation because they are essential todesert ecosystems and harbor unexplored biodiversity.

DESERT STREAMS: AN INNOVATIVE NEW APPROACH TO THEIRINVENTORY AND MAPPING by Carolyn Chainey-Davis ............................... 20A new science-based method for mapping California’s episodic desert streams incor-porates an understanding of their unique physical forms and processes where othermethods have failed.

THE ECOLOGICAL IMPORTANCE OF WASHES TODESERT BIRDS by Andrea Jones and Garry George ....... 24Audubon Society staff share a brief summary of the importance of perennial desertstreams and microphyll woodlands to migratory songbirds and other bird species inSouthern California.

JOHN O. SAWYER, JR.: 1939–2012 by James P. Smith, Jr. ..........................25

BOOK REVIEWS ........................................................................................27

2 F R E M O N T I A V O L . 4 2 , N O . 1 , J A N U A R Y 2 0 1 4

EDITORIALby Kara A. Moore

trients cycle. Deserts are rich and di-verse landscapes in which we can ob-serve the great drama of nature’s cycles.They invite adventurous explorationby our city- and computer-trained eyesas they meet the wild horizon.

On my first trip across the Mojaveand Sonoran Deserts from forestedMichigan, I was struck by the heat andby the expansive landscapes and mes-merized by the seemingly empty yetcolorful mountains, dry lake beds, andsweeping plateaus. Later, when I re-turned as a researcher exploring onfoot, I had the gift of time to see closely,to witness change over the course ofdays and seasons.

During my first desert researchproject at Ash Meadows National Wild-life Refuge, I saw the desert unfold. Iwitnessed its dynamic plants and pol-linators, lightening quick leopard liz-ards and charismatically slow deserttortoises, azure pools and pupfish,monarch and sphinx caterpillars, rag-ing flash floods, and moving soils. Oneach visit, be it to the lava fields aroundPisgah Crater or the bajadas descend-ing from Clark Mountain, I continueto find more diversity, more abundantlife, and more interesting questionsthan on the one before. Deserts chal-lenge us—as they do all of their inhab-itants—to be patient, to look closely,to build tolerance and resilience.

With this issue of Fremontia andthe next, we challenge readers to lookclosely at a region that is undergoingradical physical and landscape levelchange. In this issue, we first highlightthe diversity and ecology of the Cali-

fornia deserts. Then we focus on theincredible taxonomic diversity ofplants in general and on the biology ofrare plant species. We also emphasizethe impact that periodic water has inshaping desert ecosystems.

The second Mojave and SonoranDeserts issue, slated for distribution inMay of 2014, will cover a broad rangeof management questions. These in-clude a strong focus on the ecologicalconsequences of natural and man-made disturbances. As decisions arerapidly made regarding resource use,we need to challenge each other tocross institutional boundaries andshare information including, but notlimited to, species occurrences, popu-lation dynamics, and management fail-ures and successes. To that effect, thesecond desert issue will include ar-ticles from conservation and restora-tion practitioners and from participantsin policy debates.

There are many additional topicsof desert ecology not included in thesetwo issues. Should these issues ofFremontia peek your curiosity, Cali-fornia Deserts: An Ecological Rediscov-ery, written by long-time CNPS mem-ber and contributor Bruce Pavlik, is anexcellent read for scientists and non-scientists alike, and covers topics ofhistory, ecology, and management. Butbetter yet, visit the deserts yourself,and visit more than once. Try to seethe deserts in every season. You arelikely to gain a new, and deeper appre-ciation for these rich desert ecosys-tems, and for the great diversity of lifeforms that they support.

his is a momentous time to focusthe lens of Fremontia on theMojave and Sonoran Deserts. We

teeter on the brink of vast changes inthe North American deserts as weexpand our cities, roads, recreationalactivities, and energy generation intoone of the largest remaining frontierson the continent. Policymakers andland managers are embroiled in par-ticular on debate over the siting ofutility scale solar and wind renewableenergy facilities.

Despite the fantastic ability ofdesert ecosystems to withstand drought,endure floods, and capitalize on fa-vorable conditions when they occur,what they cannot recover from—atleast not on the scale of human lifetimes—are disturbances so intense andsizeable that they interrupt entirelythe flow of soil, water, and speciesover the landscape. The next two is-sues of Fremontia focus on the Mojaveand Sonoran Deserts and are designedto raise awareness among the public,policymakers, and renewable energyproponents of these threats and makethe strong case that deserts are uniqueand precious ecosystems that warrantvigilant protection.

The common misconception is thatdeserts are hostile wastelands, that theyare useless open spaces that may becarelessly managed. Though they haveintense climates, deserts are anythingbut barren. Even in the most extremeplaces and times, such as those iconicand seemingly barren landscapes ofDeath Valley, highly specialized insectsand plants persist, soils move, and nu-

T

The first winter storm of the season on Clark Mountain, October 2013. Hidden beyond the mountain lies Ivanpah Solar ElectricGenerating Facility, covering over 4,000 acres of former desert wilderness. It is expected to power approximately 140,000 homes whenit goes on the grid later this year. Photograph by Kara A. Moore.


FLORISTIC DIVERSITY AND DISCOVERY IN THECALIFORNIA DESERT

by James M. André

o other state in the UScan rival California’s flo-ristic diversity. The Cali-fornia Floristic Province,

which makes up the western two-thirds or cismontane portion of thestate, is celebrated widely among bi-ologists for its unparalleled diversityand high degree of endemism, andmuch of California’s extraordinarydiversity is attributed to this uniqueregion. This review, however, willfocus on the other floristic provincesof California that comprise the state’sarid transmontane deserts, a regionoften overshadowed (if not also rain-shadowed) by the California Floris-tic Province, yet of equal significanceand appeal to botanists.

The California desert, as definedby The Jepson Desert Manual(Baldwin et al. 2002), comprises theeastern third of the state and in-cludes the Great Basin Province eastof the Sierra Nevada crest and thewestern components of the Mojaveand Sonoran Deserts (Figure 1, page3). This vast region is more thantwice the size of the Sierra Nevada,spanning 500 miles from theSweetwater and White Mountainsof Mono County, southward throughDeath Valley National Park, MojaveNational Preserve, and Joshua TreeNational Park, to the lower Colo-rado Desert of San Diego and Impe-rial Counties. Consisting mostly offederal public lands, largely unpopu-lated and still unfragmented by de-velopment, the California desert isof global significance, as it repre-sents perhaps the largest intact eco-system in the US outside of Alaska.

EXCEPTIONAL DIVERSITY

Although deserts in general arecommonly portrayed in literature and

and coastal redwood trees. It is alsowhere we find California’s shortest-lived vascular plants, such as theephemeral summer annuals that cangerminate and produce viable seedin just three weeks. These same seedsmay then lie dormant in the soil formany decades before germinatingagain. And to underscore the qual-ity of the California desert ecosys-tem, naturalized alien species makeup only 8% of the flora, comparedto greater than 20% for the rest ofthe California flora.

A FLORISTIC FRONTIER

With more than 100 majormountain ranges, myriad canyons,

Source: Patrick McIntyre, UC Berkeley, 2013, modi-fied from The Jepson Manual, 2nd edition.

FIGURE 1. THE CALIFORNIA DESERTREGION AND ITS MAJOR COMPONENTS.

N lore as barren and lifeless places, theflora of the California desert is in factextraordinarily diverse. At present,approximately 2,450 native vascularplant taxa (henceforth termed “spe-cies” in this article) have been docu-mented in the California desert, rep-resenting 38% of the native speciesin the state. If the California desertregion was a US state, its flora wouldrank an impressive 18th among statesin total native vascular plant diver-sity. Even when compared to the re-nowned species-rich cismontaneCalifornia, the desert fares quite well.Mid-elevation alluvial fans of the east-ern Mojave Desert support perhapsthe highest shrub diversity in all ofCalifornia, and similar overall spe-cies diversity (90–120 totalspecies per hectare) to that ofthe primeval coastal redwoodforests of northwest Califor-nia. The remarkable richnessof our desert flora is owed inpart to its exceptional geologicand topographic diversity, aswell as the rapid speciationand diversification of largegenera (e.g., Phacelia, Astraga-lus, Eriogonum, Cryptantha,Gilia).

Though often described ashomogeneous, the desert re-gion is in fact a place of greatvariation and extremes. It iswhere the hottest and coldesttemperatures in the state arerecorded, and contains thelowest and nearly the highestelevations, and certainly thedriest climate. The desert iswhere we find the oldest vas-cular plants in California,such as creosote bush (Larreatridentata) and bristleconepine (Pinus longaeva), moreancient than the giant sequoia


playas, alkali meadows, badlandsand sprawling sand dune complexes,the California desert remains rela-tively unexplored. Yet there is a con-tinuing misconception that we have

completed our floristic inventory ofthe California desert, and that theremaining hotbeds for taxonomicdiscovery are limited to places likeIndonesia and the Brazilian Ama-

TOP: The Castle Peaks region of the Mojave National Preserve, a rich flora only recentlydocumented by botanists. • ABOVE: Early spring bloom of brittlebush (Encelia farinosa) onthe eastern slopes of the unexplored Dead Mountains near Needles, California. Allphotographs by the author.


zon. This perspective is not entirelysurprising given that in the easternUS, and even much of the west in-cluding cismontane California, fieldexploration and collecting has beenfairly comprehensive over the pasttwo centuries.

To gain a better understandingof how well we have documentedthe desert flora, I conducted anonline search of herbarium voucherrecords, sorted by local physio-graphic areas (mountain ranges,valley basins, etc.). To date, mostdesert ranges and basins have fewerthan 250 herbarium records in theConsortium of California Herbaria.And some ranges such as the Dead,Hackberry, Resting Spring, Ship,Owlshead, and West RiversideMountains, have fewer than 100records. Several major ranges, in-cluding the Turtle, Castle, Sacra-

mento, and Chemeheuvi Mountains,remained virtually uncollected un-til very recently. There are no recordsfrom the southern half of the OldWoman Mountains, one of the larg-est ranges in the eastern MojaveDesert. Although parts of the Kla-math Mountains, southern SierraNevada, and Modoc Plateau are simi-larly unexplored botanically, noother region in the state can matchthe scale of the California desert as afloristic frontier in need of docu-mentation.

Several other important obser-vations stand out when viewing thecollections history for the Califor-nia desert. Efforts to inventory anddocument desert plants, as measuredby the number of vouchers collectedper decade, have actually declinedsince the first two-thirds of the 20thcentury when famous botanists such

as Marcus E. Jones, Willis Jepson,Phillip Munz, and Robert Thorneexplored the region extensively.Though there has been a recent in-crease in general collecting by sev-eral botanists, much of our her-barium collections were made morethan 50 years ago.

The vast majority of records inthe desert occur along the relativelyfew paved roads in the region. Andwhile the more inaccessible areasremain major spatial voids for docu-mentation, temporal voids exist aswell. Summer annuals and fall-flow-ering plants are prominent in theCalifornia desert, comprising about40% of the flora, but poorly docu-mented because a disproportionatenumber of collections have beenmade in spring and early summer.Major information gaps still exist,and I make the case here that we

Sunrise on the impressive volcanic spires of the northern Turtle Mountains, a region unexplored by botanists until this past decade.


Late summer bloom of cinchweed (Pectis papposa) in the Granite Mountains of eastern San Bernardino County, showcasing the often-overlooked floristic diversity of autumn in the desert.

have only scratched the surface inour understanding of the desert flora.

What can we expect to add to theCalifornia desert flora with furtherexploration? Recent floristic studiesdemonstrate what can be accom-plished with robust and focused in-ventory and assessment. These ef-forts include, but are not limited to

the San Diego County Plant AtlasProject (desert portion) that beganin 2003 (Rebman et al.), a flora ofJoshua Tree National Park (La Doux),a flora of Mojave National Preserveand surroundings (André), and floraprojects by botanists from RanchoSanta Ana Botanic Garden (e.g,Honer, Fraga, Bell, and De Groot).

Until recently, the WhippleMountains of southeastern San Ber-nardino County represented one ofthe many botanical black holes inthe California desert. Sarah De Grootcollected the 500-square-mile rangeextensively in the mid-2000s as partof her graduate research, and docu-mented approximately 400 species,


a four-fold expansion of the flora(De Groot 2007). In the process ofadding many common species, shediscovered several new species inthe state and the California desertflora, and documented numeroussignificant range extensions.

In 2003 Joshua Tree NationalPark (JTNP) launched a compre-hensive effort to inventory the park’s1,200 square miles. Prior to this ef-fort, many areas in the park such asthe Pinto Basin, Long Canyon, andthe Coxcomb and Eagle Mountainsremained virtually unexplored. Ledby park botanist Tasha La Doux, thegoal of this project was to build uponprevious efforts, with an emphasison surveying the under-exploredspatial and temporal gaps. Over thepast 10 years the inventory at JTNPhas added 140 species to the parklist, expanding the list by 24% to itscurrent 725 species. Roughly 90%of the additions were made via fielddiscoveries, including five new spe-cies to science, while 10% wereadded through herbarium searchesand taxonomic revisions.

The final case study presentedhere is my own work on a flora ofthe Mojave National Preserve (MNP)and surrounding areas (>3,000square miles), initiated in 1995. Un-like most of the California desert,the MNP region received consider-able attention from early botanists.In particular, the Clark, New York,Providence, and Granite Moun-tains—the highest ranges in the Pre-serve—have attracted a number ofprominent collectors over the past150 years. Robert Thorne, BarryPrigge, and James Henrickson ex-plored parts of the MNP extensivelyabout 35–40 years ago, and pub-lished a flora that included the KelsoDunes and the high ranges.

In 1995 I mapped the approxi-mately 15,000 historic vouchersknown from the region at the timeand noticed that nearly all were clus-tered in the Kelso Dunes and fourhigh ranges. There were 16 othermountain ranges, such as the Cima

Cinder Cones, Castle Peaks, PiuteRange, and Woods Mountains,where few to no historic vouchersexisted. Over the course of theproject I focused approximately9,000 field hours upon these voids,and also continued to survey thehigher ranges and Kelso Dunes(Thorne et al. 1981). Since that time,120 native species have been addedto the flora (a 14% increase), manyof which were found in the “well-studied” areas (e.g., New York andProvidence Mountains). Seven ofthese species are new to science,more than 800 new rare plant oc-currences have been added, andnoteworthy range extensions con-tinue to be frequently documented.

Assuming the three flora projectsoutlined above are representative ofthe undocumented status of the en-tire California desert, we can takehome one clear message: the desertremains a floristic frontier, and theseare not the end of days for taxo-nomic discovery. On the contrary, Iwould suggest that we are in the midstof a second golden age of discovery.

RECENT DISCOVERIES ANDADDITIONS

To estimate the rate of additionsof native plant species to the Cali-fornia desert, I utilized the popularfloras that have been published overthe past 90 years. The numbers ofdesert species included in each pub-lished flora are as follows:

1925 Jepson CA Flora: 2,1491973 Munz & Supplement: 2,187

TOP TO BOTTOM: Examples of desert speciesnew to science. A close-up of the uniqueflower of the rare shrub Orocopia Moun-tains spurge (Euphorbia jaegeri). • Theshowy Clark Mountain monardella (Mon-ardella eremicola), one of several newspecies of this genus that have recently beenadded to the desert flora. • The showyClark Mountain green-gentian (Fraseraalbomarginata var. induta), only found inCalifornia on the limestone substrates ofClark Mountain.


1993 Jepson Manual: 2,2672012 Jepson Manual, 2nd ed: 2,4302013 (Present): 2,450

While there has been an increaseof approximately 300 species over90 years, representing a 14% in-crease, more than 60% of these wereadded in just the last two decades.The lull in the mid-1900s coincidedwith a national lumping trend inplant taxonomy that occurred dur-ing that period. Most of the addi-tions are newly described species,but some represent species previ-ously known only from adjacentstates or bioregions of California. Inthe past two decades the desert florahas expanded by 183 species or 8%.Over the same period, the Califor-nia flora (including the desert) hasgrown by only 5%, suggesting thatthe California desert is indeed a ma-jor hotbed for taxonomic discov-ery. In addition, I am aware of atleast 40 proposed new desert spe-cies likely to be added to the florathis decade.

This resurgent golden age of dis-covery is by no means as high as thelate 1700s and 1800s when the firstwave of exploration occurred, butit’s happening now during a timewhen field collecting (and fundingfor taxonomy) has seen a generaldecline. The current taxonomic ten-dency to split species, combined withthe fact that more people are nowdescribing new species than had beenthe case 50 years ago (and doing somore sensibly I believe) explainssome of this recent increase in num-bers of species in the Californiadesert. Tools such as Google Earthand the availability of online data-bases such as that of the Consortiumof California Herbaria also greatlyimprove the efficiency of fieldwork,especially when targeting remote ar-eas that are difficult to access.

WHAT THE FUTURE HOLDS

Based upon fairly conservativeassumptions, we can expect another

190–200 native species to be addedto the California desert flora by theyear 2100. This projection assumeswe have reached the peak rate ofdiscovery (33 species per decade),and applies a 10% decline in thenumber of additions per decade go-ing forward. Another way of look-ing at this projection is to considerwhat it means to our present levelof understanding. If 200 species areto be added to the California desertby the end of this century, and per-haps another 50–100 more in thenext century, then we can safelyassume that at least 10% of the florais presently undescribed. This isquite humbling.

The projection of 200 new addi-tions this century also assumes thatthe concept of a species remains rela-tively unchanged and that extinc-tion is not a factor (i.e., that thedesert will remain sufficiently pro-tected into the future). As to thelatter point, approximately 1,500square miles of the California desertare currently proposed for indus-trial-scale solar and wind energy de-velopment in the next several years,mostly on undisturbed federal lands.Considering indirect impacts of de-velopment (e.g., increased vectorsfor alien species invasions, losses tobiological soil crusts, barriers to dis-persal, etc.), the actual degradationof habitat will extend far beyond thefootprint of the projects themselves.According to most extinction mod-els, at this unprecedented scale andpace of impact, extinction events willbe likely.

My hope is that we continuethis push to inventory the Califor-nia desert, as we still have muchwork to do, and time is now of theessence. Perhaps more important isthat the desert gains the level ofappreciation, advocacy, and protec-tion that it deserves. To both grizzledand nascent students of botany, theCalifornia desert holds a wealth ofresearch and teaching opportuni-ties. And to those interested in pre-serving one of our last big wild

places on earth, this is your call toaction.

REFERENCES

André, J.M. 2006. A vascular flora ofthe Granite Mountains, Eastern SanBernardino County: An annotatedchecklist. Crossosoma 32(2): 38–74.

André, J.M. 2005. Inventory of Vascu-lar Plants at Mojave National Pre-serve: A Preliminary Report. Tech-nical Report #2280201178, Inven-tory & Monitoring Program, USDepartment of the Interior, NationalPark Service, Barstow, CA.

Baldwin, B.G., et al., eds. 2012. TheJepson Manual: Vascular Plants ofCalifornia. UC Press, Berkeley, CA.

Baldwin, B.G, et al., eds. 2002. TheJepson Desert Manual. UC Press, Ber-keley, California.

Barbour, M.G., and J. Major. 1988. Ter-restrial Vegetation of California. Cali-fornia Native Plant Society, Davis,CA.

De Groot, S.J. 2007. The “nose” of Cali-fornia: An important part of thestate’s diversity. Fremontia Vol. 35(No 1): 3–6

Hickman, J.C. ed. 1993. The JepsonManual of Higher Plants of Califor-nia. UC Press, Berkeley, CA.

Jepson Flora Project. 2013. JepsoneFlora, http://ucjeps.berkeley.edu.

Jepson, W.L. 1925. A Manual of theFlowering Plants of California. UCPress. Berkeley, CA.

McLaughlin, S.P. 1995. Floristic rela-tionships of the eastern MojaveDesert: A quantitative analysis of lo-cal floras. Crossosoma 21:57–74.

Munz, P.A. 1959. A Flora of California.UC Press, Berkeley, CA.

Stein, B.A. 2002. States of the Union:Ranking America’s Biodiversity. ANatureserve Report. The NatureConservancy, Arlington, VA.

Thorne, R.F., B.A. Prigge, and J.Henrickson. 1981. A flora of thehigher ranges and the Kelso dunesof the Eastern Mojave Desert in Cali-fornia. Aliso 10(1): 71–186.

James M. André, University of CaliforniaRiverside, Department of Biology, UCNRSGranite Mountains Desert ResearchCenter, HC1 Box 101, Kelso, CA 92309,[email protected]


RARE PLANT DIVERSITY IN THE CALIFORNIA DESERTS:PRIORITIES FOR RESEARCH AND CONSERVATION

Kara A. Moore and James M. André

mong people of all culturesthere is an inherent ap-preciation of things thatare rare and unique,

whether a precious gemstone orgreen flash on the horizon at sunset.To the conservation scientist, rareorganisms are of particular interestbecause they are genetically distinctaspects of living diversity that maybe at risk of extinction, or may sig-nify an impaired ecosystem. As stew-ards of the planet, most of us believethat we have an ethical obligation topreserve biodiversity and the eco-logical processes that sustain natu-ral systems. This goal is especiallychallenging, given that our under-standing of how healthy ecosystems

function and the role that each lifeform plays is incomplete. The uniqueand diverse array of rare plants inthe California deserts presents sucha conservation challenge because thedistributions and basic life historiesof most species remain poorly docu-mented.

While endangered mammals andbirds often command more publicattention, plants form the basis ofall terrestrial life by providing en-ergy, materials, and habitat struc-ture. Globally, 25–40% of native vas-cular plant taxa (henceforth termed“species” in this article) are at risk(Pitman 2002). Within NorthAmerica, numbers of imperiledplants may be most underestimated

in our deserts because they havesystematically received far less at-tention from botanists than otherbioregions. In the California deserts,native plants are being added to theflora faster than any other region ofthe state. For more on this excep-tional floristic frontier, see the otherarticle by James M. André on page 3of this issue.

The California Native PlantSociety’s (CNPS) Inventory of Rareand Endangered Plants ranks 35%of California’s native vascular floraas being of conservation concern,but only 20% of the California desertflora is similarly ranked. There areat least three reasons for this lowerpercentage. First, endemism (where

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California bearpoppy (Arctomecon merriamii), a rare perennial found in small populations on calcareous gravel deposits of the northeasternMojave Desert and in western Nevada. Photograph by Erica Smith.

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a species is only found in a restrictedhabitat or location) is lower in theCalifornia deserts than in the rest ofCalifornia. Yet even though the Cali-fornia deserts represent only a frac-tion of the Great Basin, Mojave, andSonoran Deserts, a striking 25% ofthe rare species in the Californiadeserts are endemic to the state,highlighting the uniqueness of theCalifornia desert flora within theNorth American deserts. Second,because the deserts are, at present,relatively pristine compared to muchof California, most rare desert plantsare inherently rare, with only a hand-ful driven to rarity by human activi-ties. Third, we have underestimatedthe actual number of rare plant spe-cies in the California deserts—par-

ticularly given that the region is ahotbed for new species discovery—so most new additions to the florawill be rare.

The California deserts are un-dergoing rapid change as a result ofincreasing use by humans, especiallyon federal public lands targeted forwidespread solar and wind energydevelopment. Planning measures forrare plant protection will be basedupon limited existing knowledge ofthe desert flora, and thus lack thenecessary level of detail. Even afterwe document the distributions ofrare plants, we are challenged tounderstand their unique biology.

In this article we focus on keyfactors that influence rarity in theCalifornia desert flora and provide a

basis for establishing strategies forconservation. Our goal is to bringattention to the diversity of raredesert plants and to provide insighton their ecology that will be usefulto those planning for their protec-tion. We use case studies to illus-trate some of the unique facets ofrare desert plants that influence theirmanagement, highlighting addi-tional research needs and priorities.

OPPORTUNISTS BYNECESSITY

Desert plants must be adaptedto a harsh and variable climate topersist. These adaptations includethe climatic responsiveness of an-

Fantastic bloom in the foreground of the rare Bartow wooly sunflower (Eriophyllum mohavese)—a plant that is often less than a centimeterin width and height—in 2011 near Kramer Junction in the northwestern Mojave Desert of California. In infrequent favorable years, thesetiny rare sunflowers form an unusually dense carpet under the common annual Bigelow’s tickseed (Coreopsis bigelovii). Photograph byKara Moore. Close-up photograph by James André.

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nuals, the seasonality of herbaceousand woody perennials, and the en-during architecture of many succu-lents and shrubs. Desert annualsarise from a long-lived dormant seedbank only when triggered by theright combination of temperatureand moisture. Seed dormancy buff-ers annual populations from climaticfluctuations by permitting emer-gence only when conditions arelikely to lead to high survival (Claussand Venable 2000). Many desertannuals have a very limited periodduring which germination and emer-gence can occur, and it is highlydependent upon precipitation. Insome cases, especially in the sum-mer, this window may be only a fewweeks. The absence of abovegroundplants over multiple years can cre-ate the false impression that plantpopulations are no longer present,although they are merely lying inwait as viable seed in the soil forfavorable germination conditions.One plant that demonstrates this dra-matic year-to-year variation is theminiscule Barstow wooly sunflower(Eriophyllum mohavense), a rare win-ter annual found only near KramerJunction and Edwards Air Force Base(photo on page 10).

The long dormancy periods ofrare herbaceous desert perennialsmust be considered in the assess-ment of population densities and indeveloping long-term monitoringprograms. The flowering phenologyof some herbaceous perennials canbe highly complex. For example,scarlet four o’clock (Mirabilis cocci-nea) typically flowers in response tolocalized summer precipitation, butmay also respond to late winter rainsand then flower in early summer.Some deciduous shrubs such as veradulce (Aloysia wrightii) and desertmilkwort (Polygala acanthoclada)remain leafless until they receivesufficient precipitation. Many mon-soon-responsive summer- and fall-flowering desert species such asdesert purslane (Portulaca hali-moides) apply a different photosyn-

thetic pathway (C4) thanthe majority of plant spe-cies (C3), which allowsthem to more efficientlygrow and reproduce in thiswarm climate with sparseand/or unpredictable rain-fall.

The challenge of limitedfield documentation andperiodic emergence is ex-emplified by Mojave milk-weed (Asclepias nycta-ginifolia), a tuber-formingperennial limited to theeastern Mojave. Prior to sur-veys for the construction ofthe Ivanpah Solar Elec-tric Generating System(ISEGS)—a massive projectthat has been built on ap-proximately 3,740 acres offederal public land—themilkweed was known inCalifornia from only fouroccurrences, all above 3,000feet in elevation, and all atsome distance from theproject site. Mojave milk-weed was thus assumed tohave low potential to occurin Ivanpah Valley and wasoverlooked during initial project sur-veys, which were conducted duringfairly dry years. Follow-up surveysin subsequent years of increasedrainfall revealed numerous popula-tions in the ISEGS footprint. Theseplants had been dormant and unac-counted for during the preliminaryenvironmental review process.

RARE PLANT HABITATS

The causes of rarity in the desertare nearly as diverse as the rare floraitself (e.g. Rabinowitz 1981; Fielder1986, 2001). Many rare desert spe-cies are narrow endemics that occurin a single restricted geographic area,and/or may be confined to a uniquehabitat to which they have adapted.They may have either recentlyevolved in place (neoendemics), orbe ancient species (paleoendemics)

that have become restricted and nowpersist in an isolated location suchas a mountaintop. Unique deserthabitats rich in endemic rare spe-cies include characteristic uplands

Desert milkwort (Polygala acanthoclada)in bloom. Photographs by James André.


such as sand dunes, gypsum claydeposits, and limestone substrates.Deserts also include localized wet-land ecosystems that are home tomany rare plants, including alkalimeadows, non-alkali wetlands,freshwater springs, riparian systems,and infrequently inundated land-scape features such as playas andwashes (Pavlik 2008).

Another challenge for rare habi-tat specialists is the limited extent,fragmentation, and disjunction ofmany of their specialized habitats.For example, species specialized ongypsum clay (gypsophiles) and lime-stone substrates such as Californiabearpoppy (Arctomecon merriamii)have populations scattered through-out suitable habitat patches that canbe separated by many miles. Lime-stone ranges that extend into Cali-fornia from Nevada, such as ClarkMountain and the Nopah Range,house a suite of limestone special-ists that are rare in California, butfairly common in Nevada wherelimestone substrates are abundant.

Rare species may also be distrib-uted across a range of ubiquitous

habitats, their distribution and rar-ity being driven by other factors.These include a long list of taxasuch as Orocopia Mountains spurge(Euphorbia jaegeri), Mojave mon-keyflower (Mimulus mohavensis),and the crucifixion thorn (Castelaemoryi). These are fragmented ordisjunct populations that are diffi-cult to explain ecologically. Causesof rarity for such species might in-clude geographic isolation due tolong-term climate change, alteredpollinator guilds, restricted geneflow, or other factors. Additionalresearch is often required to discernwhether such widely scattered popu-lations are genetically isolated acrossthe range of the species’ distribu-tion.

INFLUENCES OFREPRODUCTIVE BIOLOGY

In developing conservation strat-egies for rare desert plants it is of cri-tical importance to first understandtheir diverse reproductive systems.

Of the many factors that can in-

fluence rarity, the production andfates of seeds are often the mostcritical and least understood, andthe complex interaction of flowersand fruits with animals warrants ex-tensive research. Some rare speciesemploy self-incompatibility, mean-ing that individuals need pollen fromanother individual to produce vi-able seed. These species, such asLittle San Bernardino Mountainslinanthus (Linanthus maculatus),need larger population sizes to per-sist than species that readily self-pollinate.

Idiosyncrasies of rare plants in-clude atypical chromosome num-bers, low genetic diversity, and rela-tionships with specialist pollinatorsor dispersers. And these factors com-monly compound to influence rarespecies. For example, Peirson’s milk-vetch (Astragalus magdalenae var.peirsonii) is restricted to the sandsof the Algodones Dunes, is self-in-compatible, and requires pollinationby a single native bee species (Groomet al. 2007). In addition, its habitathas been severely degraded by off-road-vehicle use. Even when seedsare produced, their ability to sur-vive, disperse, and persist in the seedbank is not assured. Rare plants atAsh Meadows and Pisgah Crater canlose up to 90% of their total seedproduction due to herbivory by jack-rabbits, sometimes for several con-secutive years (Pavlik et al. 2009).Rapid construction of large-scaleenergy projects in the Californiadesert could exacerbate seed lossesdue to local distortion of food webs.

NEEDS AND PRIORITIES

We must put our feet to the earth.There is an ongoing need for inven-tory and mapping of known rareplant species in the California desertsto better inform regional conserva-tion and management. The majorityof rare plant records occur within amile of major roads, in historicallypopular botanical areas (e.g., NewYork Mountains, Mecca Hills), or

A PROFILE OF CALIFORNIA’S RARE DESERT FLORA

he 491 listed rare California desert species are an exceptionallydiverse group with representatives in 70 vascular plant families.

Nearly a quarter of these species (152) are either endemic to theCalifornia deserts or rare both here and elsewhere, placing them inthe highest rarity class, California Rare Plant Rank (CRPR) 1B. Alarge proportion (209 species, 43%) are CRPR 2B, meaning they arerare in California but more common in other states. This is a rela-tively high percentage of 2B species in comparison to the rest ofCalifornia. Another 118 plants are listed as CRPR 4, a watch list ofspecies with limited distributions.

Of the CNPS-listed desert species, only 13 have federal listingstatus (are protected) under the Endangered Species Act, with 8receiving protection as “Endangered” and the other 5 listed as “Threat-ened.” Federal listing of desert plants is often sidelined by othercritical research needs or because officials may assume that rarespecies occurring on federal lands are sufficiently protected. Yet wehave identified an additional 20–30 desert species that potentiallymeet criteria for federal listing status, and listing would greatlyimprove their prospects for research funding and protection.

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in areas surveyed prior to develop-ment. Geographic models can beused in some cases to estimate spe-cies ranges, but have important limi-tations when applied to rare andunder-surveyed species (see articleby Patrick McIntyre on page 15).Given that the California desertsremain a floristic frontier (we haveadded 183 species in the past twodecades alone), extensive inventoryis still needed to catalog additions tothe native flora. Many undiscoveredspecies will also be rare, and notafforded the necessary protectionsuntil their taxonomy is confirmedand their distributions mapped. Acomprehensive desert-wide inven-tory will take institutional will, muchexpertise, and many decades to com-plete.

In addition, research on the ecol-ogy of California’s rare desert spe-cies is vital for stewarding popula-tions amid rapidly increasing threats,

such as from urbanization, utility-scale energy, mining, recreationalactivities, invasive species, and cli-mate change. At present, status re-ports that document basic biologyand perceived threats are availablefor only about a quarter of the 491known rare desert plants, and fewerthan 5% have accompanying conser-vation management plans. Of equalconcern is the fact that establishedlong-term monitoring or researchprograms exist for less than 1% of allrare plants in the California desert.We need to increase and extend spe-cies-specific research to provide abaseline understanding of a repre-sentative set of rare desert species.

Basic population studies that fol-low the fates of individual plantsand their progeny are especiallyneeded to make predictions aboutthe effects of landscape fragmenta-tion, population reduction, and cli-mate change on the long-term per-

sistence of species. A comprehen-sive life history approach that in-cludes monitoring of all life stages,including seed banks, is necessary inorder to make realistic predictionson population viability. Each stageof growth and reproduction can begreatly affected by other species inthe community, some negatively(competition and herbivory) andsome positively (pollination, seeddispersal). Therefore it is necessaryto implement research that includesinteracting species and is conductedover several decades. Such a timeframe can better capture populationresponses to fluctuations in climateand variations in species interactions.Unfortunately, such long-term stud-ies are extremely rare.

Currently, we are assembling 13years of field data and observationson white-margined beardtongue(Penstemon albomarginatus), an her-baceous perennial known in Cali-

White-margined beardtongue (Penstemon albomarginatus) in bloom near the Sleeping Beauty Mountains, Central Mojave Desert, California.Despite the numerous flowers in 2011, extremely few fruits and seeds were successfully produced because of strong herbivory and otherunknown limits on fruit development. Photograph by Kara Moore.


fornia from a single population nearPisgah Crater. We are assessing howdifferent life stages, precipitation,and species interactions affect itsability to persist. We found thatcaging plants from black-tailedjackrabbit herbivory increases sur-vival and seed production, but per-haps not enough to protect the spe-cies from other factors that threatenlocal extinction. Increased droughtfrequency, as predicted by global cli-mate change models, radically in-creases the probability of extinction.Additional demographic monitoringis needed to determine if this popu-lation will rebound or rapidly ad-vance to California Rare Plant Rank1A status (plants presumed extinctin the state).

For many desert rare plants, in-cluding the white-margined beard-tongue, perhaps the most curiousand important life stage is the mostchallenging to study: the soil seedbank. Unknown factors include thelongevity of seeds under field con-ditions, mechanisms and rates of dis-persal, local density, and conditionsthat trigger germination. Seedlingemergence, determined by environ-mental factors controlling germina-tion and mortality in the soil, is of-ten the most critical driver of popu-lation growth. When mathematicalmodels are used to simulate popula-tions of white-margined beard-tongue, small variations in the mea-sured rates of seedling emergencecan result in very different proba-bilities of population growth andpersistence.

We also must improve our un-derstanding of rare species’ interac-tions with other plants, pollinators,mutualists, and herbivores. Manyplants endure strong herbivory, atleast periodically, when populationsof insects, jackrabbits, pocket go-phers, wood rats, or other speciesreach high densities (often cyclic ordue to a lack of predators), or dur-ing drought when food resourcesare limited. For example, in someyears, alkali mariposa lily (Calo-

chortus striatus) and crowned muilla(Muilla coronata) can be nearly im-possible to observe before herbivoresconsume aboveground leaves andstems. However, for the majority ofrare species, by far the most criticalinteractions are those they have di-rectly and indirectly with humans.

A CALL FOR RESEARCH

Ongoing research is needed todetermine the diversity, abundance,and distributions of rare desert plantsand to understand the fundamen-tals of their ecology. We emphasizethat the most critical facet of inven-tory and mapping for rare species isto remember that absence cannot beassumed from a single survey (oreven a few) for species that have adormant life phase. And with rapidand extensive changes in arid landuse, we need to advance taxonomicresearch on both known species andthose that are not yet described inorder to forward their protection.

There is a pressing need for popu-lation biology research on rare plantspecies. Focal points must includeresearch on the detection and func-tion of dormant life stages and thepositive and negative effects of spe-cies interactions. We must deepenour understanding of how both dor-mant life stages and interactions withother species vary with fluctuationsin climate that are characteristic ofthese ecosystems. Furthermore, re-search to explore the effects of prox-imity to development, landscapefragmentation, soil erosion, alteredhydrology, and other anthropogenicdisturbances on rare plants is criti-cal to their persistence.

Regional and global processessuch as atmospheric nitrogen depo-sition and global climate change havefar-reaching effects on local popula-tions and must be included in thescope of studies. Although species-specific management is required bythe biological uniqueness of rare spe-cies, clearly all forms of life are bestserved by protecting functioning

ecosystem and landscape units. Pres-ervation of large contiguous and di-verse wildlands throughout the Cali-fornia deserts is the only way toensure protection of its many rarespecies.

REFERENCES

Clauss, M.J., and D.L. Venable. 2000.Seed germination in desert annuals:An empirical test of adaptive bethedging. American Naturalist 155:168–186.

Fielder, P.L. 1986. Concepts of rarityin vascular plant species, with spe-cial reference to the genus Calo-chortus Pursh (Liliaceae). Taxon 35:502–518.

Fielder, P.L. CNPS Inventory, 6th Edi-tion, 2001.

Groom, J.D., et al. 2007. Quantifyingoff-highway vehicle impacts on den-sity and survival of a threateneddune-endemic plant. Biological Con-servation 135(1): 119–134.

IUCN 2013. The IUCN Red List ofThreatened Species. Version 2013.1.http://www.iucnredlist.org.

Pavlik, B.M. 2008. The CaliforniaDeserts: An Ecological Rediscovery.University of California Press, Ber-keley, CA.

Pavlik, B.M., Moore-O’Leary, K.A., andA.E. Stanton. 2009. Quantifying her-bivore impacts on rare plants at AshMeadows. Proceedings of the Califor-nia Native Plant Society ConservationConference.

Pitman, C.A., and P.M. Jørgensen.2002. Estimating the size of theworld’s threatened flora. Science (1November 2002): 989.

Rabinowitz, D. 1981.Seven forms ofrarity. In The Biological Aspects ofRare Plant Conservation, ed. H.Synge. John Wiley & Sons Ltd., NewYork, NY.

Kara A. Moore, Department of Evolutionand Ecology, University of California,One Shields Avenue, Davis, CA 95616,[email protected]; James M.André, Department of Biology, Universityof California, Riverside, and UC NaturalReserve System, Granite Mountains DesertResearch Center, HC1 Box 101, Kelso,CA 92309, [email protected]


APPLYING SPECIES DISTRIBUTION MODELINGTO IDENTIFY RARE SPECIES HOT SPOTS

by Patrick McIntyre

n the last 15 years, ecologists havedeveloped mapping tools to pre-dict new areas where rare speciesmay occur, based on their known

localities. These tools are broadlyreferred to as species distribution orecological niche models. (Ecologistsdebate whether the method predictsthe ecological niche of a species ormerely where the species occurs onthe landscape.) Species distributionmodeling identifies aspects of theenvironment, such as elevation, tem-perature, and rainfall associated withwhere a species is known to occur.

Using mathematical models andGeographic Information System(GIS) technology, ecologists can

then make a map of predicted habi-tat for a species beyond areas whereit is currently known by identifyingplaces with similar habitat. Com-bining these tools with predictedchanges in temperature and rainfall,ecologists can also make predictionsabout where species have the poten-tial to occur in the future, given dif-ferent scenarios of climate change.

A MAP OF RARE PLANTHOTSPOTS

To illustrate how these tools caninform plant conservation, I builtspecies distribution models for 151

rare plant species whose docu-mented occurences allowed us toidentify at least 5 populations. Using

ITwo California endemics known from theOrd Mountains southeast of Barstow, apredicted hotspot of rare plant diversity.LEFT: Mohave monkeyflower (Mimulusmohavensis). • RIGHT: Creamy blazing star(Mentzelia tridentata). Both photographsby the author.

LEFT: MAP OF CRPR 1 PLANTS (PRIMARILY CALIFORNIA ENDEMICS). • RIGHT: MAP OF RARE OR THREATENED PLANTS INCALIFORNIA AND WHICH OCCUR MORE COMMONLY OUTSIDE OF THE STATE.

PREDICTIVE MAPS OF RARE PLANT DIVERSITY IN THE CALIFORNIA DESERTS.

Source: Patrick McIntyre, UC Berkeley. Predictive mapping of rare desert plants, 2013.

Note: Areas in warmer colors (yellows and browns) are areas of higher predicted rare plant diversity. The predicted region correspondsto the proposed boundaries of the Desert Regional Energy Conservation Plan.


a standard set of climate data for the20th century (worldclim.org), Imade maps of where these plantswere predicted to occur across theDesert Renewable Energy Conser-vation Plan (DRECP) area. This areais the recent focus of rapid develop-ment and conservation planning. Ithen overlaid predictions for Cali-fornia Rare Plant Rank (CRPR) 1species (those primarily endemicto California) and CRPR 2 species(those rare in California but morecommon elsewhere) in order to iden-tify predicted regions of rare plantdiversity in the California deserts(see maps on page 15).

Several patterns emerge from ex-amination of the maps of predicteddiversity. First, western areas of thedesert abutting mountain ranges areidentified as areas of high diversity.This highlights how these areas sup-port both true desert species and

species from other habitats that ex-tend to the edge of California’sMojave and Sonoran Deserts. Sec-ondly, the Clark and Kingstonmountain ranges in eastern Califor-nia, much of which are included inthe Mojave National Preserve, areidentified as centers of diversity bothfor endemic and non-endemic taxa.The Preserve’s ranges host many spe-cies that normally occur outside ofCalifornia, but also occur in thesemountains.

Another intriguing area of pre-dicted high diversity is the Ord andLavabed mountain ranges southeastof Barstow. This area falls outsideof National Park/National Preserveareas and currently is known tosupport a number of rare Califor-nia endemics, such as the creamyblazing star (Mentzelia tridentata),Mohave monkey flower (Mimulusmohavensis), and Mojave menodora

(Menodora spinescens var. moha-vensis), and is predicted to have suit-able conditions for several other rarespecies. Based on the simple mapspresented here, this area might sup-port additional rare species not cur-rently documented in those loca-tions, or might be a region that couldact as a refuge for plants from otherareas forced to shift their ranges as aresult of climate change.

Predictive maps such as the onesaccompanying this article representvaluable tools that can help guidefield-based efforts to document plantdiversity. They can identify unan-ticipated locations where rare spe-cies might be found. They can alsobe used to help predict where ap-propriate habitat for rare plantscould occur over large areas that areinfeasible to survey by foot. In addi-tion, they provide a means for pre-dicting where species might occurin the future under scenarios of cli-mate change, something that can-not be accomplished through fieldsurveys.

However, predictive maps arebased on imperfect data—known lo-calities that represent only a subsetof each species’ real distribution.They are not substitutes for on-the-ground exploration by experiencedbotanists. It’s one thing to use a spe-cies distribution model to identifylikely areas to hunt for new rareplant occurrences, and another thingentirely to use the predictions of amodel to decide which parcels ofland to preserve and which to de-velop. Finally, models can’t predictthe distribution of a species that hasnever been described, and new spe-cies are described every year in theCalifornia desert. These models areuseful tools that we can use to guideresearch and focus field explorationbased on what we know today, andwhat might likely occur in the fu-ture.

Patrick McIntyre, University of Califor-nia, 3101 Valley Life Sciences, Berkeley,CA 94720, [email protected]

Four rare plants known from the Clark and Kingston ranges in the eastern Mojave desertof California, a known and modeled hotspot of rare plant diversity in California. CLOCKWISE

FROM TOP LEFT: Stephen’s beardtongue (Penstemon stephensii). Photograph by Jim André. •Mohave milkweed (Asclepias nyctaginifolia). Photograph by Drew Maraglia. • Parish’s clubcholla (Grusonia parishii). Photograph by Kara Moore. • Mojave desert plum (Prunuseremophila). Photograph by Jim André.


THE CASTLE MOUNTAINS:A HOTSPOT OF DESERT PLANT DIVERSITY

by Duncan Bell

he Castle Mountains are afantastic example of the greatdiversity of relatively smallmountain ranges in the Cali-

fornia deserts. In San BernardinoCounty, the Castle Mountains areonly approximately 30 square miles,yet they are home to over 30 rareplant species and hundreds of com-mon species. Enter any canyon ofthe Castle Mountains, scramble ontoany ridgeline, or walk any wash andyou will find unique and interestingplant species, some of which arefound only in this rugged corner ofthe Mojave, such as canyon bird’sfoot (Lotus argyraeus var. multi-caulis) and the showy pinto beard-tongue (Penstemon bicolor).

The Castle Mountains, publiclyowned by the Bureau of Land Man-agement, are best accessed by HartMine Road from the Mojave NationalPreserve. Access requires a highclearance vehicle, and four-wheeldrive is seasonally necessary. The

center of the Castle Mountains isrugged and rocky, with hidden can-yons containing a diversity of raredesert annuals, including nine-awned pappus grass (Enneapogondesvauxii) and Clark Mountainspurge (Euphorbia exstipulata).

Steep canyons spill out into widevalleys that surround the range. Theyare home to dense and extraordinar-ily healthy stands of Joshua trees thatare part of a desert savannah con-taining a diversity of native annualand perennial grasses. Around twodozen grass species are found here,of which half a dozen are rare grasses.Some species, such as burro grass(Scleropogon brevifolius) and falsebuffalo grass (Munroa squarrosa), arepart of unique desert grasslandsfound nowhere else in California.Also present are many other rare plantpopulations such as matted cholla(Grusonia parishii), Abert’s sanvitaliaaster (Sanvitalia abertii), and red fouro’clock (Mirabilis coccinea).

The botanical frontier of theCastle Mountains has only recentlybeen breached by botanists suchas James André, Andy Sanders, andmyself. Every year we are findingnew and previously unknown rareplant populations, and certainlymore discoveries remain. For ex-ample, in 2012 both James Andréand I found Mexican panicgrass(Panicum hirticaule) in the CastleMountains area, the first collectionsof this species for San BernardinoCounty and an 88-mile northernrange extension.

Every field press that is filledwith care in the Castle Mountains islikely to hold important discoveriesthat will allow us to further under-stand our rare plant populations andtheir distribution in the rugged butfragile California deserts.

Duncan S. Bell, 1500 North College Ave-nue, Claremont, CA 91711-3157, [email protected]

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Joshua tree woodlands amidst a unique assemblage of grasses at sunset in a valley surrounding the Castle Mountains. Photograph by theauthor.


MICROBIOTIC SOIL CRUST COMMUNITIES:A CRITICAL COMPONENT OF CALIFORNIA’S DESERTS

by Nicole Pietrasiak and Jeffrey R. Johansen

ow more than ever, pris-tine and fragile desertecosystems are facingmajor threats from hu-

man development, and efforts arebeing made to protect sensitive, spe-cial-status plant and animal species.In contrast, microbial communitiesof desert soils have been neglectedin conservation efforts, although theyare often considered as critical toecosystem health. It is well docu-mented that the microbial commu-nities within biological soil crusts

are especially crucial to the ecologi-cal functioning of desert ecosystems.The crusts harbor diverse taxa in-cluding mosses, lichens, fungi, greenalgae, diatoms, and cyanobacteriathat bind together mineral soil par-ticles into water- and wind-stableaggregates at the soil surface. Thiscrust is vital because it prevents ero-sion in sparsely vegetated landscapes.

In addition to their importantrole in stabilizing soil surfaces, theircontribution to soil fertility is essen-tial. Some crust microbes are capable

of converting atmospheric nitrogento ammonium, an essential but lim-ited nutrient in desert systems. Thus,crusts represent an important nitro-gen source for associated vascularplant communities or soil food webs.Biological soil crusts also can seques-ter substantial amounts of carbon.The carbon can accumulate in themicrobial biomass, ultimately add-ing soil organic matter to the sys-tem. This organic matter is then avail-able in the soil food web. In fact,biological soil crusts are rightfully

Biological soil crusts of Californian Deserts. • BELOW: The darkened soil surface indicates the presence of lichen and algal crusts. Themeasuring tape gives an idea of scale. • FACING PAGE, TOP: An algal crust is all that holds together soil particles on this desert slope, whichcontains little vegetation. • FACING PAGE, BOTTOM: A picture taken with a microscope of a potential new cyanobacterial genus and speciesfound in a wilderness area of Joshua Tree National Park. All photographs by Nicole Pietrasiak.

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considered to be the fertile mantle ofdesert landscapes.

The physiological performanceof crust communities and the re-sulting ecological impact in the land-scape varies with community com-position. Depending on climatic,geomorphic, and soil properties,crust communities differ in the struc-tural components of microbioticcrusts (lichens, mosses, fungi,cyanobacteria). Where this happens,crust communities have distinctforms and structures. The Great Ba-sin, Mojave, and Sonoran Desertsare especially rich in crust types,

which vary with regardsto the dominance ofmosses, lichens, algae,and fungi. In the MojaveDesert, significant differ-ences in soil aggregate sta-bility, nitrogen fixation,and carbon fixation canbe detected among crusttypes. For example, li-chen and moss crust com-munities contribute mostto these ecosystem func-tions, whereas someweakly-developed algalcrusts perform poorly.Hence, the effect crustshave on an area dependsboth on the communitytypes present and howmuch of the ground iscovered.

In addition to the im-portant ecological role ofthese communities, theyharbor a multitude of un-discovered species andgenera. A research teamled by Valerie Fletchnerinvestigated over 100strains of eukaryotic al-gae and discovered 15clades (a clade is a groupof organisms whose mem-bers share features from acommon ancestor), ofwhich only 6 could beidentified to establishedgenera. Recently, Radka

Muehlsteinova, together with theauthors, investigated over 150 cy-anobacterial strains and found atleast 23 clades, with most of theselineages being potential new generaand species to science. These find-ings are just an initial indication ofhow little we know about thebiodiversity of these amazing mi-croscopic communities.

In summary, one of the greatestanthropogenic threats to desert sys-tems is the loss of biological soilcrust communities, and the con-comitant loss of biodiversity, ero-sion control, and soil fertility.

REFERENCES

Belnap, J., and O.L. Lange. 2003. Bio-logical Soil Crusts: Structure, Func-tion, and Management, 2nd ed.Springer Verlag, Berlin.

Evans, R.D., and J.R. Johansen. 1999.Microbiotic crusts and ecosystemprocesses. Critical Reviews in PlantSciences 18: 183–225.

Flechtner, V.R., N. Pietrasiak, and L.A.Lewis. 2013. Newly revealed diver-sity of eukaryotic algae from wilder-ness areas of Joshua Tree NationalPark (JTNP). Monographs of theWestern North American Naturalist6:43–63.

Garcia–Pichel, F., et al. 2003. Small-scale vertical distribution of bacte-rial biomass and diversity in biologi-cal soil crusts from arid lands in theColorado Plateau. Microbial Ecology–46: 312–321.

Herrick, J.E., et al. 2010. Fine gravelcontrols hydrologic and erodibilityresponses to trampling disturbancefor coarse-textured soils with weakcyanobacterial crusts. Catena 83:119–126.

Lovich, J.E., and D. Bainbridge. 1999.Anthropogenic degradation of theSouthernCalifornia desert ecosys-tem and prospects for natural recov-ery and restoration. EnvironmentalManagement 24: 309–326.

Pietrasiak, N., J.R. Johansen, and R.E.Drenovsky. 2011. Geologic compo-sition influences distribution of mi-crobiotic crusts in the Mojave andColorado Deserts at the regionalscale. Soil Biology and Biochemistry43: 967–974.

Pietrasiak, N., et al. 2011. Spatial dis-tribution and comparison of distur-bance impacts to microbiotic soilcrust in the Little San BernardinoMountains of Joshua Tree NationalPark, California. Western NorthAmerican Naturalist 71: 539–552.

Pietrasiak, N., et al. 2013. Biologicalsoil crust community types differ inkey ecological functions. Soil Biol-ogy and Biochemistry 65: 168–171.

Nicole Pietrasiak and Jeffrey R. Johansen,Biology Department, John Carroll Uni-versity, 1 John Carroll Boulevard, Univer-sity Heights, OH 44118, [email protected], [email protected]


DESERT STREAMS: AN INNOVATIVE NEW APPROACHTO THEIR INVENTORY AND MAPPING

by Carolyn Chainey-Davis

One of the most startling paradoxesof the world’s drylands is that al-though they are lands of little rain,the details of their surfaces aremostly the products of the actionof rivers. To understand the natu-ral environments of drylands is tounderstand the process and formsof their rivers.

—W.L. Graf (1988)

A PARADOX AND APARADIGM SHIFT

he physical processes thatshape streams in the desertare very different from thosein other parts of California.

The hydrology of desert streamshas been likened to an early descrip-tion of life in the trenches of WorldWar I: “long periods of boredom,brief moments of terror.” Desert

streams—driven by wildly unpre-dictable rain patterns, heavy floodflows, and sparsely vegetated anderodible soils—have a form and hy-drology that are remarkably differ-ent from streams elsewhere in thestate.

Concepts and methods used todelineate streams that were devel-oped for streams in areas of greaterrainfall, and an historical bias forperennial streams, has led to an un-derestimation of the extent and im-portance of desert streams. With theincreasing pressure to develop re-newable energy projects in thedesert, there is an urgent need foran entirely new approach to the waywe perceive, map, inventory, andevaluate desert streams, one that rec-ognizes and incorporates theirunique physical, hydrological, andecological processes. A new science-based geomorphic and ecological

method for delineating episodicstreams in the California desert suc-cessfully meets that challenge whereother methods have failed.

Surprisingly, one of the mostprominent features of desert land-scapes is their system of stream chan-nels (Schwinning et al. 2011). In-deed, sometimes entire alluvialfans—those vast triangular apronsof sand, gravel, and arid soils thatare built from arid mountain can-yons and comprise 30–40% of thedesert landscape—are characterizedby complex networks of activelyshifting stream channels. These wa-ter-driven landscape features breathelife into otherwise harsh environ-ments, but are routinely ignored orundervalued in development plan-ning and environmental impact as-sessments.

Until recently, change cameslowly to the California desert. Habi-

T

Although smaller streams may not always support classic desert wash plants, the denser and more robust vegetation along even smallerstreams provides shade, escape cover, breeding and nesting sites, seed, and other food sources. Unnamed distributary channel on desertpavement near El Centro, CA. Photograph by Kris Vyverberg, California Department of Fish and Wildlife.


tats were generally intact or mini-mally impacted, with disturbancegenerally limited to highways, rail-road and utility corridors, scatteredmining operations, and sheep graz-ing. Populations of many of thedesert’s rare plants and animals wereconsidered relatively stable. But inthe last decade the political and eco-nomic push for renewable energydevelopment has placed many desertspecies and ecosystems at risk, eco-systems inextricably linked to thefunctional integrity of streams andtheir floodplains.

ECOSYSTEM BENEFITS

All streams are life giving, butperhaps nowhere more so than indeserts. Streams occur in a varietyof channel forms in the Mojave andSonoran Deserts, but there are veryfew perennial streams (those thatflow year-round). The most com-mon are the episodic streams, whichinclude ephemeral streams (flowingbriefly during and after a rain event)and intermittent streams (those thatflow seasonally). Although they look

and behave differently thanperennial streams, episodicstreams provide all of thesame ecosystem services—ecological, hydrological,and physical (Levick et al.2008). However, our cul-tural bias for perennialstreams has led to a dis-missal of the desert’s epi-sodic streams and theunique habitats they sup-port as having little or nohabitat value.

The fundamental im-portance of ephemeral andintermittent streams towildlife in the desert is un-disputed. Streams and theirfloodplains provide notonly critical wildlife habi-tat, but provide a founda-tion for much of the desert’s bioticdiversity. Higher soil moisture con-tent and topographic relief of desertstreams provide shade and coolertemperatures, which are used ex-tensively by wildlife, and especiallyby less mobile creatures that cannotavoid the harsh environment out-side the stream ecosystem. Thehigher moisture content, gravel,sand, and soils delivered by streamflows, and the rapid drainage prop-erties of these loose, stream-depos-ited sediments support plant com-munities that are distinct from thoseon upland soils. This contrast is par-ticularly stark on desert pavementswhere in some areas nearly all thewildlife habitat is found alongstreams, even in the smallest ofchannels.

The streamside vegetation oflarger ephemeral streams can bequite distinct: the floriferous wood-lands of palo verde (Parkinsoniafloridum), ironwood (Olneya teso-ta), and smoketree (Psorothamnusspinosus) that characterize the largecompound channels of the SonoranDesert region of California; or theoases of lush Fremont cottonwood(Populus fremontii) and sandbar wil-low (Salix exigua) on wide reaches

of the Mojave River; or the distinc-tive shrubby thickets of catclaw(Senegalia greggii) or desert almond(Prunus fasciculata) on the single-thread channels of desert mountaincanyons.

In comparison to larger streams,the smaller channels of alluvial fansand their broad active floodplainsmay not be as floristically distinctfrom adjacent plant communities.Yet they are nearly always measur-ably denser, more robust, and havegreater vegetative cover than theadjacent drier uplands. The smallerchannels play an equally importantrole in seed dispersal and germina-tion, including the seeds of somerare species. The hard seed coats ofsome species, especially the legumi-nous plants like ironwood, paloverde, catclaw, and smoketree, areabraded by the flows, promotingmore efficient germination in seedsthat may otherwise require years ofweathering to germinate.

Where small streams dissipateon the valley floor, water sometimespools at the terminus of the chan-nel, and provides valuable tempo-rary access to water for wildlife.Pooled water and fine sediments cangerminate abundant annuals follow-

Distributary channel networks on alluvial fans runningoff the Avawatz Mountains near Death Valley, CA.They provide groundwater recharge and storm runoffto playa lakes and replenish the sand source criticalfor the maintenance of dunes. Oblique view of GoogleEarth imagery provided by Jeremy Lancaster, CaliforniaGeological Survey.

Pooled water at the terminus of a smallchannel where it dissipates near the toe ofan alluvial fan, three days after a quarter-inch summer storm event. Two weeks later,the channel was filled with native summerannuals. Pahrump Valley, CA. Photographby Carolyn Chainey-Davis.


ing summer storms, like Bigelow’smonkeyflower (Mimulus bigelovii),scarlet lupine (Lupinus concinnus),and shaggyfruit pepperweed (Lepi-dium lasiocarpum). These providefresh young greens for habitat andforage in a dry summer landscape.

Reptiles and amphibians arepresent in higher diversity in desertstreams than uplands, and manyspecies preferentially use streamsand their floodplains because ofdenser plant cover (Baxter 1988).Desert streams provide important

food sources for desert tortoise andother species because they supportmore diverse plant communitieswith greater cover and higher diver-sity of summer and fall annuals. Theyalso host greater bird abundance anddiversity, in fact up to 1.5 timesmore breeding species and twice asmany wintering species than thedrier adjacent desert scrub (Kubickand Remsen 1977; Tomoff 1974;Daniels and Boyd 1979a, 1979b).

The ecosystem benefits provid-ed by desert streams extend wellbeyond watercourse boundaries.Downstream and downwind duneecosystems are wholly dependent onthe replenishment of sands deliv-ered by episodic streams to the“Aeolian” (wind-based) sand trans-port systems that maintain dunesand the many rare animals and plantsthey support, such as CoachellaValley milk-vetch (Astragalus len-tiginosus var. coachellae). Episodicstreams provide storm water runoffto playa lakes and recharge thegroundwater sources, which in turnsupport the unique and often rarehabitats that occur at the playa mar-gins: marshes, saltgrass meadows,alkali sink scrubs, and dune thick-ets. Where the playas and wetlandsoccur along migratory pathways,they seasonally provide critical

invertebrate resources and restingopportunities for migrating shore-birds on their journey across thedesert (Robinson and Oring 1996).

NEW METHODS TO MAPDESERT STREAMS

As pressure to develop the desertincreases, it has quickly becomeapparent that 1) the old conceptsand methods used to delineatestreams in wetter regions—whichassume regular and reliable flows,well-defined channels, or the pres-ence of classic riparian zones—don’t work in the desert; and 2)there is an urgent need to developa reliable and consistent methodfor mapping and evaluating streamsthat incorporates an understandingof the distinctive physical, hydro-logic, and ecological processes ofdesert streams.

In response, the California De-partment of Fish and Wildlife(CDFW), with additional fundingfrom the California Energy Com-mission (CEC), is preparing a sci-ence-based methodology for the de-lineation of desert streams to assistagency staff and energy project de-velopers, and to help interested par-ties in their review of project im-

Unnamed stream east of Twentynine Palms, San Bernardino County, California. The smaller swales that dissect the upper fan surfacescollect runoff and initiate stream flow from the surrounding, less permeable dark desert pavement surfaces. These are important sourcesof water, sediment, nutrients, and other materials during runoff, and are considered integral parts of a desert stream system. Photographby Jeremy Lancaster, California Geological Survey.

Dense stringers of galleta grass (Hilariarigida) line the small channels of an alluvialfan on the Palo Verde Mesa near Blythe,California, providing predator protectionand a cooler microclimate for wildlife.Photograph by Roland Brady, Brady andAssociates Geological Services.


pacts to streams. Though still in itsinfancy, MESA: Mapping EpisodicStream Activity (Vyverberg and Brady2013) provides a photographic atlasof the geomorphic indicators of epi-sodic stream activity, mapping guid-ance, guidance in the interpretationof aerial photos, and six stream map-ping case studies. Even to the non-professional this field guide, whichis filled with interesting photos anddiagrams, provides a riveting crashcourse in desert fluvial geomorph-ology (the shaping of rivers andstreams). This document will beavailable on the CEC and CDFWwebsites by spring 2014.

STREAM INFLUENCE FARBEYOND THE CHANNEL

Stream channels are among themost conspicuous features on thedesert landscape and contribute tothe ecological health of desert plantcommunities (Schwinning et al.2011). Thus, when we modify oreliminate desert streams we essen-tially halt life-giving flow to anentire watershed. For example,multiple studies that compare thevegetation of unaltered alluvial fansto the dewatered portions down-stream of a diversion have foundcompelling results. No matter howsmall the channels nor the frequencyor duration of flows, the health andintegrity of alluvial fan ecosys-tems depends on the integrity oftheir stream channel networks(Schwinning et al. 2011).

Ecological impacts and the im-pacts on stream-forming processesare suffered far beyond the originaldisturbance sites (Schwinning et al.2011; Schlesinger and Jones 1984;Ludwig 1986; Johnson et al. 1975).In one study near Joshua Tree Na-tional Park (Schlesinger and Jones1984) that compared vegetation ontransects of 4,350–4,750 feet aboveand below a railroad that bisects analluvial fan, creosote bush suffered a50% reduction in biomass on the

dewatered portions of the fan, andshrub density was reduced over 30%.Such studies provide an importantnew perspective on the function,complex structure, and intrinsicvalue of distributary channel net-works, regardless of their size or thefrequency of their flows. The desertis an enchanting place, home to manyremarkable plants and animals thatare wonders of adaptation. The en-during health of these species andtheir living habitats are undeniablylinked to the functional integrity ofthe desert’s complex drainage sys-tems. The challenge for the conser-vation, management, and restorationof the desert’s unique episodic streamecosystems begins with our percep-tion of them. They are highly dy-namic systems in time and space andtheir influence, particularly on allu-vial fans, spans the entire watershed.

Regardless of the duration oftheir flows, episodic streams pro-vide all the same ecosystem benefitsof perennial streams, and their eco-logical importance is particularlysignificant given their vast extent.MESA: Mapping Episodic Stream Ac-tivity by Vyverberg and Brady (2013)is an important starting point in thedevelopment of a new science-basedmethod for mapping, inventorying,and evaluating desert streams thatincorporates an understanding oftheir unique forms and processes.

REFERENCES

Baxter, R.J. 2008. Spatial distributionof desert tortoises (Gopherus agassi-zii) at Twentynine Palms, California:Implications for relocations. In Man-agement of Amphibians, Reptiles, andSmall Mammals in North America.Proceedings of the Symposium, July19–21, 1988. US Forest Service Gen-eral Technical Report RM-166. Flag-staff, AZ.

Daniels, B.E., and J. Boyd. 1979a.Forty-second breeding bird census.No. 142. Ironwood-smoketree desertwash. American Birds 33(1): 92–93.

Daniels, B.E., and J. Boyd. 1979b.Forty-second breeding bird census.

No. 150. Paloverde desert wash.American Birds 33(1): 94.

Graf, W.L. 1988. Fluvial Processes inDryland Rivers. Springer-Verlag, Ber-lin, Germany.

Johnson, H.B, F.C. Vasek, and T. Yon-kers. 1975. Productivity, diversity,and stability relationships in MojaveDesert roadside vegetation. Bulletinof the Torrey Botanical Club 102: 116–115.

Kubik, M.R., Jr., and J.V. Remsen, Jr.1977. Fortieth breeding bird census.No. 125. Catclaw-rabbitbrush desertwash. American Birds 31(1): 75–76.

Levick, L., et al. 2008. The Ecologicaland Hydrological Significance ofEphemeral and Intermittent Streamsin the Arid and Semi-Arid AmericanSouthwest. U.S. Environmental Pro-tection Agency and USDA/ARSSouthwest Watershed ResearchCenter. EPA/600/R-08/134, ARS/233046. Washington, DC.

Ludwig, J.A. 1986. Primary produc-tion variability i5n desert ecosys-tems. In Pattern and Process in DesertEcosystems, ed. W.G. Whitford. Uni-versity of New Mexico Press, Albu-querque, NM.

Robinson, J.A., and L.W. Oring. 1996.Long-distance movement by Ameri-can avocets and black-necked stilts.Journal of Field Ornithology 67: 307–320.

Schlesinger, W.H., and C.S. Jones.1984. The comparative importanceof overland runoff and mean annualrainfall to shrub communities of theMojave Desert. Botanical Gazette145: 116–124.

Schwinning, S., et al. 2011. The influ-ence of stream5 channels on distri-butions of Larrea tridentata and Am-brosia dumosa in the Mojave Desert,CA, USA: Patterns, mechanisms, andeffects of stream redistribution.Ecohydrology 4(1): 12–25.

Tomoff, C.S. 1974. Avian species diver-sity in desert scrub. Ecology 55: 396–403.

Vyverberg, K.A., and R. Brady. 2013.MESA: Mapping Episodic StreamActivity. California Department ofFish and Wildlife. September 2013draft (unpublished). Sacramento,CA.

Carolyn Chainey-Davis, 182 Grove Street,Nevada City, CA 95959 [email protected]


THE ECOLOGICAL IMPORTANCE OF WASHESTO DESERT BIRDS

by Andrea Jones and Garry George

esert riparian habitats inSouthern California arerare and of additional con-servation concern because

they represent critical habitat for en-dangered breeding bird species suchas Southwestern Willow Flycatch-ers, Western Yellow-Billed Cuckoos,and Least Bell’s Vireos. In addition,in vast desert regions, these greenpatches with fresh water supportthousands of songbird migrantswhile they are en route from theirwintering grounds in Central andSouth America to breeding groundsfurther north.

Of particular note to avian com-munities are desert washes—drystreambeds subject to rapid flowduring flash flooding—in the Colo-rado and Mojave Deserts, which oc-cur in both upland and riparian habi-tat, and may include trees, shrubs,or a combination of both. In A Natu-ral History of the Sonoran Desert

(2000), Mark Dimmitt says that “drywashes occupy less than 5% of thissubsection of the Sonoran Desert,but support 90% of its bird life.” Thestructural diversity in desert washescan include trees such as palo verde(Parkinsonia aculeata) and ironwood(Olneya tesota) that provide breed-ing and feeding habitat for a varietyof songbirds, particularly Phaino-peplas, Ash-Throated Flycatchers,Verdin, Crissal, LeConte’s and Ben-dire’s Thrashers, Long-Eared andWestern Screech Owls, Black-TailedGnatcatchers, Gila and Ladder-Backed Woodpeckers, Lucy’s War-blers, Northern Mockingbirds, andLoggerhead Shrikes, some of whichare California bird species of specialconcern.

The National Audubon Societyhas recognized the significance ofthese habitats by naming the “Colo-rado Desert Microphyll Woodland”an Important Bird Area. This area

includes significant stands of mi-crophyll woodland associated withdesert washes that flow east intothe Colorado River, including theChemehuevi Wash, Vidal Wash, andMiltiptas Wash. Another major oc-currence of microphyll woodlandhabitat lies at the base of theAlgodones Dunes, halfway betweenthe Colorado River and the south-ern end of the Salton Sea. In spring,these areas attract large concentra-tions of migratory songbirds andprovide nesting areas for breedingspecies such as the Elf Owl, GilaWoodpecker, Long-Eared Owls, andBendire’s Thrashers. Additional in-formation is available at: http://www.prbo.org/calpif/htmldocs/desert.htm.

Andrea Jones, Suite 14, 601 Embarcadero,Morro Bay, CA 93442, [email protected]; Garry George, 926 S. Citrus Ave, LosAngeles, CA 90036, [email protected]

D

Desert riparian areas are relatively rare, yet provide critical breeding and migratory habitat for many endangered bird species. Photographby Andrea Jones.

INSETS: (TOP) An endangered desert bird, the Least Bell’s Vireo, depends on desert washes and riparian habitats to breed. Photograph bySteve Mlodinow. • (MIDDLE) The Verdin, distinguished by its yellow facial pattern, is often found in desert areas containing desert washes.Photograph by Greg Smith. • (BOTTOM) Le Conte’s Thrasher is a bird that breeds in desert scrub and desert wash habitat in SouthernCalifornia, including the Carrizo Plain. Photograph by Alan Schmierer.


JOHN O. SAWYER, JR.: 1939–2012by James P. Smith, Jr.

ohn Sawyer passed away at hishome in Arcata on August 19,2012. His loss is deeply felt byhis wife, Jane Cole, his family,

and his many colleagues and friendswith whom he had close professionaland personal relationships. We willmiss his comradeship, infectiouslaugh and smile, support, boundlessenergy, and wealth of knowledge.With his passing, our state has lostone of its most respected and influ-ential plant ecologists. The Califor-nia Native Plant Society has lost aformer president, an effective advo-cate for the preservation of our na-tive flora, and an individual who con-tinued to make significant contribu-tions to the society for over 40 years.

John earned his bachelor’s degreeat California State University, Chico,and his MS and PhD in plant ecologyfrom Purdue University. While pur-suing his graduate degrees, he wasan instructor at CSU, Chico, a gradu-ate assistant at Purdue, and a fieldecologist for a consulting firm withprojects in Costa Rica and Thailand.

Sawyer joined the faculty ofHumboldt State University in 1966where he remained until his retire-ment in 2001. He taught general ecol-ogy, plant ecology, arctic and alpineecology, California natural history,and plant taxonomy lab sections.Early on, he was the prime mover ina very popular spring break offer-ing—the desert field trip—an inten-sive excursion in a state bus fromArcata to Southern California. In allof these classes John demonstratedcritical thinking, and he insisted onit in his students as well. He wasespecially proud of his cadre of about50 graduate students, many of whomremained close friends and who nowoccupy professorships and respon-sible positions around the country.

John’s favorite instructionalarena was the field, where he ex-

celled. It was there that students orparticipants in CNPS or Jepson Her-barium field trips experienced hispassion and detailed knowledge ofplants. He was not known for lead-ing casual strolls across the coun-tryside. The lack of a well-definedtrail was only a slight impediment ifthere was a manzanita or a goose-berry to be examined.

John’s mobility became increas-ingly limited by the effects of thecancer that would take his life. Soon our private sojourns together,we just changed our mode of opera-tion to day trips in the car. Johnwould spot a plant, usually of thewoody persuasion, we would screechto a stop, and Jane and I wouldgather specimens for him to look at.

His countless hours in the field,extending well beyond his formalretirement, resulted in more than40 books and papers! There wereexcursions into such diverse topicsas the biological formations of theeastern United States, the vegeta-tion life zones of Costa Rica, andeven a flora of a region in northernThailand. But John’s firstpassion was the vegeta-tion of California, par-ticularly that of thenorthwestern part of thestate. Having gone intothe field with him for 43years, I can attest to bothhis boundless enthusi-asm and incredibleknowledge.

In particular, as hisstudents would say, Johnhad this thing for woodyplants, especially coni-fers. You can see this soclearly in his book,Northwest California: ANatural History. He wasalso a coauthor, alongwith his close colleague

Todd Keeler-Wolf, and Julie Evens, of the secondedition of A Manual of California Veg-etation. It is recognized as the stan-dard reference on the subject.

John wrote sections for Terres-trial Vegetation of California on themontane and subalpine vegetationof the Klamath Mountains, the for-ests of northwestern California, andalpine vegetation. His Trees andShrubs of California, coauthored withJohn Stuart, a Humboldt State col-league, has become a favorite fieldguide. He also produced a book withAndrea Pickart, a former graduatestudent, on the ecology and restora-tion of coastal dunes in NorthernCalifornia.

Some plant ecologists have littleknowledge of the flora itself. Not sowith John. He was involved withDale Thornburgh in documenting arange extension of subalpine fir intoCalifornia; authored a paper on theserpentine flora of the Lassics; co-authored the description of a newsubspecies of Frangula; and authoredor coauthored four family treatments

J

John out on the Bridge of Heaven trail in Colorado’s SanJuan Mountains, July 2001. Photograph by Todd Keeler-Wolf.

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in The Jepson Manual and the treat-ment of Rhamnaceae that will ap-pear in the Flora of North AmericaNorth of Mexico.

John and I wrote 5 editions ofour Keys to the Families and Generaand Vascular Plants in NorthwesternCalifornia, and 22 editions of ourchecklist of the plants of that sameregion have appeared. We were work-ing on a specimen-based checklist ofthe vascular plants of the KlamathRegion, and Sawyer exacted a prom-ise that I would see it completed.

John became the fourth presi-dent of CNPS in 1973, the first “out-sider” to occupy the position, in thesense of not being one of the groupthat founded the society. He beganthe process of making CNPS a state-wide organization. Board meetingsthat had been held on Thursday eve-nings, typically in the home of a BayArea member, were moved to Satur-day. An unfinished set of bylawsthat had not permitted chapter presi-dents to vote at board meetings wasrewritten. He would later serve as amember of five state-level commit-tees: Conservation, EducationalGrants, Publications, Rare Plants,and Vegetation. I doubt that anyoneelse can match that record of suchdiverse committee service.

He was an editor of the secondedition of the CNPS Inventory of Rareand Endangered Vascular Plants of

California. John was a frequent con-tributor to the CNPS Bulletin andFremontia. His first article, appear-ing in the CNPS Newsletter in 1973(later to become Fremontia), was onthe Klamath Region. A bibliographyof John’s publications will soon beavailable at the Humboldt State Uni-versity Herbarium website (www.humboldt.edu/herbarium/).

John was a founding member ofthe North Coast Chapter of CNPS in1970 and served as its first president.He was especially interested in thevegetation and flora of the Klamathregion of northwestern California andsouthwestern Oregon. His field tripsto the Lanphere Dunes, Russian Peak,The Trinity Alps, Scott Mountain,Stony Creek Bog, Walker Ridge, andserpentine areas were legendary.

His local conservation effortswere exceptional. John was a criti-cal player in the establishment ofthe Lanphere Dunes near Arcata andin maintaining its natural state byorganizing “lupine bashes” to eradi-cate the invasive yellow bush lupine(Lupinus arboreus). The coastaldunes were also the home of one hismajor research efforts on the statusof the Menzies’ wallflower (Erysi-mum menziesii).

He was also intimately involvedin the work that led to the designa-tion of the Russian Wilderness inSiskiyou County. There he and DaleThornburgh would document theoccurrence of 17 conifers in a one-square-mile area around Little DuckLake, which must make it one of therichest conifer forests in the world.John and Jane assisted our friendand colleague, Dave Imper, in pro-tecting serpentine barrens in TheLassics (Humboldt and Trinity coun-ties), the home of the endemicLassics lupine (Lupinus constancei).

Among his many honors, Johnwas recognized by Humboldt StateUniversity as its 1997 Scholar of theYear, by the California Botanical So-ciety in the 2008 dedication of itsjournal, Madroño, and by CNPS inmaking him a Fellow in 1994. In

2011 he received the J.C. PritzlaffConservation Award from the SantaBarbara Botanical Garden. He wasactive in several organizations, in-cluding as a Councillor of the Savethe Redwoods League and in theEcological Society of America, wherehe served as a technical advisor.

In addition to this impressivelist of accomplishments, John wasalso an avid hiker, serious photog-rapher, world traveler, follower ofPurdue University football, habituéof art museums, student of the op-era, and an admirer of the music ofBach, Villa Lobos, Piazolla, Ives,Loreena McKennitt, Cecilia Bartoli,Bob Dylan, and Judy Collins.

John would not have wanted atraditional memorial service. In itsplace, Jane, her sister Susan, MichaelKauffmann (a fellow “conehead”),and I organized a celebration ofJohn’s life and accomplishments atPatrick’s Point State Park, just northof Arcata. About a hundred peopleattended. As you would expect, therewas a time for recollections. Ithought that I knew John pretty well,but I was amazed at stories that werenew to me, at the depth of emotionexpressed, and at how important hehad been in the lives of so manywho were there and others who sentremembrances to be read.

If I might end on another per-sonal note, someone once observedthat you are lucky in life if you haveone really good friend. If so, I wasvery lucky indeed.

Memorial contributions may bemade to any of the organizationsmentioned above, to a memorialbench that will be dedicated to Johnat the Lanphere Dunes (Friends ofthe Dunes, P.O. Box 186, Arcata95518), or to the John O. Sawyer, Jr.Endowment to support field studiesin botany (Advancement Founda-tion, Gift Processing Center,Humboldt State University, 1 HarpstStreet, Arcata 95521-8299).

James P. Smith, Jr., 884 Diamond Drive,Arcata, CA 95521, [email protected]

LEFT TO RIGHT: John Sawyer, Virginia Keeler-Wolf, and his wife Jane Cole, enjoying aride in an outboard up the Yacumama Riverof the Amazon Basin, Peru, summer 2002.Photograph by Todd Keeler-Wolf.


I Am the World’s Oldest-Known LivingTree by Dr. Gilbert G. Thibault. 2012.Winmark Communications, Glendale,AZ. 56 pp, $21.56 hardcover. ISBN#978-1-89222-515-3.

This book is dedicated “to the chil-dren of the world and their parents . . .and to the wise people who had theforesight to preserve and protect thisunique habitat. . . . ” This fine 2012children’s book about bristlecone pineswill delight all readers and deepen theirappreciation of this remote native.

From the first leaf, author/photog-rapher Gil Thibault’s graphic expertiseis evident. Soft greyed illustrations pointout detail the reader might miss in thestriking photos, giving the reader theability to see the forest and the trees. Atone of respect for these trees is set inthe beautifully detailed introductorymap. Section themes pique curiosity,particularly the bristlecone as a weatherindicator, historian, or survivor. Stun-ning photos of bristlecone pines againstthe night sky are accompanied by aninvitation to visit them, and an assur-ance that “ . . . we will leave the lights ofthe Milky Way on for you.”

A master storyteller, Thibault en-gages the reader in devising names forcreatures and trees and speculatingabout phenomena. He informs with fas-cinating facts: scientists have founddowned deadwood that’s 11,000 yearsold, and the trees do not die of old age!Humor threads through nearly everypage: “I have heard some people say myfoliage is very green for my age.” Chil-dren will love the extremes he describes,from stormy winter weather to drought.

Told from the perspective of TheMethuselah, a 5,000-year-old bristle-cone pine, the book is unabashedlyanthropomorphic. And in an interest-ing variant of personification, Methu-selah suggests humans can learn adapt-ability and resilience from bristleconepines. A fascinating four-page timelineplaces these trees amid human habita-tion, discovery, and invention.

Children of all ages will be drawnto the rich imagery the scientific lan-guage elicits, from dendrochronology,contiguous, and fortitude, to dolomite.

Thibault takes the opportunity to teachgood science rather than avoid topicscontentious to a few. There are severalreferences to the potential impact ofclimate change on the bristlecones. Theauthor notes that they are threatened insummer by bark beetles, which die offin cold weather. A rise in global tem-peratures may endanger them and ben-efit the bark beetle, since warmer tem-peratures encourage the developmentof softer wood, which bark beetles pre-fer. On the other hand, there is a missedopportunity to explore balanced eco-logical relationships: pine bark beetlesare eaten by Clark’s Nutcrackers, whichdisperse bristlecone seeds. In nearbyyellow pine forests, bark beetles are anormal part of fire ecology.

In an adaptation that improves onJoyce Kilmer’s sing-song cadence,Thibault adapts his well-known poem“Trees” to celebrate the bristlecone.Also included is a folksong, “Bristle-cone Pine,” by Hugh Prestwood, alongwith an online video link. The lyricsevoke the forest, but mentions of Jesusand heaven and hell may put off non-Christian readers. Along the same trail,the timeline would benefit from morenon-Western entries. Small criticismsin a stunning book.

While a broad audience will becaptivated by I Am the Oldest Known

Living Tree, teach-ers and parents canuse the text to meetthe goals of thestate’s curriculumstandards in scienceliteracy, particu-larly in grades 3–6.They expect students will “activelyseek the wide, deep, and thoughtfulengagement with high-quality literaryand informational texts that buildknowledge, enlarge experience, andbroaden worldviews.” This book cer-tainly qualifies as such a text.

Author Gil Thibault has written avolume that’s full of wonder, visualand verbal treasures, accurate science,humor, and reverence for bristleconesand their habitat. I Am the Oldest KnownLiving Tree makes one want to ensurethat these venerable trees survive forgenerations to come.

—Barbara Roemer andMerry Byles-Daly

BOOK REVIEWS

CORRECTIONThe photo caption on p. 2 of Fremontia,Vol. 41, No. 3 (September 2013) iden-tified Fremont Canyon as located inIrvine County. It is actually located nearthe city of Irvine, but in Orange County.


The plant guide that follows is orga-nized alphabetically by family and,within family, more loosely by genusand taxonomic relatedness. A tremen-dous amount of information has beenpacked into the book; literally no inchof space is wasted and nearly everyspecies is not only succinctly described,but also represented by multiple pho-tographs.

Even local variations within a spe-cies such as occur in California buck-wheat (Eriogonum fasciculatum), coastgoldenbush (Isocoma menziesii), andgum plant (Grindelia camporum) arediagrammed or described with photo-graphs. Key anatomical and morpho-logical traits such as floret structure insunflowers or flower keel shape anddistribution of hairs in lupines are alsoclearly diagrammed to allow for easieridentification.

Allen and Roberts’ book has sev-eral other unique and refreshing fea-tures that place it in the top tier of localnatural history references. Pollinatorsand animal guilds are described forkey species and habitats. For example,

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nine insect species associated withmilkweed and dogbane are described(with photographs!) for the reader. Theetymology of all genus and speciesnames is provided. Symbols are addedfor sensitive species that have federalor state protected status, as well as forthose of current local concern.

While the guide is extremely thor-ough, readers should note that it is notall-inclusive. A scattering of less showyspecies such as walnut, oak, alder,members of the goosefoot family, andall grasses and their relatives are ex-cluded to save space. Several non-na-tive plants occurring in wildland habi-tats are also missing. Keeping bothwould have moved the book out of therealm of a field guide and into that of adesk reference.

Most refreshing to me is that theauthors take a holistic view of plantsin our fragile landscapes. What theyhave produced will not only serve as afield guide to specialists and lay people,but also as an encyclopedia of localplants and their ecology.

—Jutta Burger

Wildflowers of Orange County and theSanta Ana Mountains by Robert L. Allenand Fred M. Roberts, Jr. 2013. LagunaWilderness Press, Laguna Beach, CA.498 pages, $35.00, soft-cover. ISBN978-0-9840007-1-5.

Southern Californiaplant enthusiasts—espe-cially those in OrangeCounty—have reason tocelebrate the publicationof a new book. Wildflow-ers of Orange County andthe Santa Ana Mountains,by Robert (Bob) Allenand Fred Roberts, is aglorious nearly 500-pageguide to the showy floraof Orange County andadjacent areas.

Nearly ten years have passed sincerumors of this ambitious project firstsurfaced, and the end product will thrillreaders. The authors are two child-hood classmates from Orange Countywho evolved along parallel paths asbotanists and natural historians. BobAllen, known to his followers as “BugBob,” is well known for his extensivelocal knowledge of plants and insects,as well as his entertaining public pre-sentations and classes. Wildflowers be-gan as his brain child. Fred Roberts isthe leading authority on status anddistribution of plants in Orange Countyand has enriched the county as botani-cal guru and artist for years. Together,they are the perfect team to have cre-ated this work.

The authors first introduce the re-gion, habitat types, and terminologyneeded to identify flowering plants.

F R E M O N T I AV O L . 4 2 , N O . 1 , J A N U A R Y 2 0 1 4

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Please make your check payable to “CNPS” and send to: California Native Plant Society, 2707 K Street, Suite 1, Sacra-mento, CA 95816-5130. Phone: (916) 447-2677; Fax: (916) 447-2727; Web site: www.cnps.org.; Email: [email protected]

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CNPS members and others areinvited to submit articles for pub-lication in Fremontia. If inter-ested, please first send a shortsummary or outline of whatyou’d like to cover in your ar-ticle to Fremontia editor, BobHass, at [email protected]. Instruc-tions for contributors can befound on the CNPS website,www.cnps.org, under Publica-tions/Fremontia.

Fremontia Editorial AdvisoryBoardSusan D’Alcamo, Jim Andre,Ellen Dean, Phyllis M. Faber,Holly Forbes, Dan Gluesenkamp,Brett Hall, David Keil, PamMuick, Bart O’Brien, RogerRaiche, Teresa Sholars, DickTurner, Mike Vasey

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FROM THE EDITOR

(continued on inside back cover)

James M. André is director of the Granite Mountains DesertResearch Center, senior advisor to the CNPS Rare PlantProgram (RPP), and chair of the RPP Committee.

Duncan Bell is a field botanist for Rancho Santa Ana BotanicGarden, and spends most of his time in the undercollectedand unexplored mountain ranges of Southern California.

Jutta Burger, PhD, is the managing director of the scienceand stewardship department at the Irvine Ranch Conservancyin Irvine, California.

Merry Byles-Daly and Barbara Roemer are members ofthe Redbud Chapter of CNPS and elementary educators.Together they have logged over 60 years in the classroom.

Carolyn Chainey-Davis is a botanical consultant whorecently conducted analyses of the biological impacts of sixproposed solar power plants in the California desert.

Garry George directs the Chapter Network and renewableenergy programs for Audubon California.

Jeffrey R. Johansen is professor of biology at John CarrollUniversity.

Andrea Jones directs the Coastal Stewardship Program forAudubon California.

Patrick McIntyre is an ecologist with the Berkeley Initia-tive in Global Change Biology at UC Berkeley. His researchincludes predicting rare desert plant distributions.

Kara A. Moore is an ecologist in the Center for PopulationBiology at UC Davis and studies the dynamics that influ-ence species distributions and the persistence of rare plants.

Nicole Pietrasiak is a post-doctoral researcher in the biol-ogy department at John Carroll University.

James P. Smith, Jr. is professor of botany, emeritus, andcurator of the herbarium at Humboldt State University inArcata, California.

his issue and the next of Fremontia, two issues focusedon California’s deserts, exemplify what CNPS doesbest. As the leading organization concerned with the

health of the state’s native plants and vegetation types, webring together in one place science-based informationgleaned from the work of our own staff and a cadre of CNPScitizen-scientists, along with that of other prominent scien-tists from academia, government agencies, and independentconsultants. This information is then available to guidethose who are entrusted to make decisions about how bestto manage and conserve our environment.

For those who know little about California’s deserts,these two issues will be eye-openers, and are likely to dispelthe myth that a desert area is a wasteland devoid of ecologi-cal value. One of the reasons some may hold that view isbecause the richness of deserts is not readily apparent to theuntrained eye. Much is hidden by nature to protect it fromthe comparative harshness of the desert ecosystem. But afterreading the articles in this issue, you may be amazed at allyou discover should you decide to take a hike in any ofCalifornia’s deserts.

We owe a debt of gratitude to those researchers whoare willing to endure the challenges presented by desertsin order to inform us of what is happening there. Research-ers are a unique, unheralded lot, and their work demandsconsiderable patience, precision, dedication, and a system-atic approach. Each adds their own piece to the jigsaw ofscientific knowledge. While it is exciting to discover newspecies and occurrences and to work in beautiful surround-ings, most botanists I know are more concerned with pro-tecting our native plants and ensuring that their vitality andbeauty endures.

—Bob Hass

T

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Documents

VOL. 42, NO. 1 • JANUARY 2014 FREMONTIA...VOL. 42, NO. 1, JANUARY 2014 FREMONTIA 1 CONTENTS THE COVER: Sunrise on the east side of the Granite Mountains in the Mojave National Preserve