CALIFORNIA'S PRAIRIES AND GRASSLANDS CALIFORNIA'S ...

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FREMONTIA VOLUME 39:2/39:3, MAY/SEPTEMBER 2011 JOURNAL OF THE CALIFORNIA NATIVE PLANT SOCIETY $10.00 (Free to Members) VOL. 39, NO. 2 AND VOL. 39, NO. 3 MAY 2011 AND SEPTEMBER 2011 FREMONTIA CALIFORNIA’S PRAIRIES AND GRASSLANDS CALIFORNIA’S PRAIRIES AND GRASSLANDS

Transcript of CALIFORNIA'S PRAIRIES AND GRASSLANDS CALIFORNIA'S ...

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JOURNAL OF THE CALIFORNIA NATIVE PLANT SOCIETY

$10.00 (Free to Members)

VOL. 39, NO. 2 AND VOL. 39, NO. 3 • MAY 2011 AND SEPTEMBER 2011

FREMONTIA

CALIFORNIA’S PRAIRIESAND GRASSLANDS

CALIFORNIA’S PRAIRIESAND GRASSLANDS

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STAFF (SACRAMENTO)Executive Director . Dan GluesenkampFinance & Administration Manager .

Cari PorterMembership & Development Coor-

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

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

EvensVegetation Ecologists . . . . . . Jennifer

Buck-Diaz, Kendra SikesEducation Program Director . . . Josie

CrawfordAdministrative Asst. . . . Marcy MillettOffice Asst. . . . . . . Caroline Garland

STAFF (AT LARGE)Fremontia and CNPS Bulletin Editor . .

Bob HassLegislative Consultant .Vern GoehringEast Bay Conservation Analyst . . . . .

Mack CastermanWebsite Coordinator . . Mark Naftzger

PROGRAM ADVISORSRare Plant Program Senior Advisor . . .

Jim AndreVegetation Program Senior Advisor . .

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

Laura CampCNPS Press Director . . . Nancy MorinPoster Program . . . Bertha McKinley,

Wilma Follette

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

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

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

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

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

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

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

Joan A. JerneganRiverside/San Bernardino counties . .

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

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

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

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

Ellen Edelson

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

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

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

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

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

VOL. 39, NO. 2, MAY 2011 AND

VOL 39, NO. 3, SEPTEMBER 2011

F R E M O N T I A

Copyright © 2011California Native Plant Society

Disclaimer:

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

CALIFORNIA NATIVEPLANT SOCIETY

Dedicated to the Preservation ofthe California Native Flora

Bob Hass, EditorGlen Holstein, Contributing Editor

Beth Hansen-Winter, Designer

Brad Jenkins and Cynthia Roye,Proofreaders

CALIFORNIA NATIVE PLANT SOCIETY

MEMBERSHIPMembership form located on inside back cover;

dues include subscriptions to Fremontia and the CNPS Bulletin

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

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CONTENTSPRAIRIES AND GRASSLANDS: WHAT’S IN A NAME? by Glen Holstein ............................................................. 2The accepted name for California’s prairie/grassland landscapes has changed back and forth and may significantlyimpact its conservation.

IS THERE STILL NATIVE DIVERSITY IN CALIFORNIA GRASSLANDS? by Megan E. Lulow andTruman P. Young ............................................................................................................................................................... 6Surveys of valley and south coastal grasslands support claims for placing a greater emphasis on native wildflowers whenreferring to the composition of native grasslands.

UNDERSTANDING CALIFORNIA GRASSLAND ECOLOGY by Paula M. Schiffman .......................................... 12Species composition of California grasslands is, to a great extent, influenced by two natural factors: variable rainfall andsoil disturbance produced by native animals.

BETWEEN THE BLADES: THE EVOLUTION OF PLANT DIVERSITY IN CALIFORNIA’S PRAIRIES ANDGRASSLANDS by Ellen Dean ....................................................................................................................................... 16Our assemblages of native prairie wildflowers have come together over millions of years of geologic time to enchant uswith their colorful displays.

GEOLOGY, CLIMATE, AND CALIFORNIA PRAIRIE DISTRIBUTION by Glen Holstein ...................................... 22Rediscovering California prairie’s lost landscapes and reexamining why they’re there.

VERTEBRATES OF CALIFORNIA GRASSLANDS by Gene R. Trapp ...................................................................... 31To what degree do California’s native vertebrates depend on grasslands for their survival? Is it complete dependence fora given species, or less than that?

INVERTEBRATES OF CALIFORNIA’S PRAIRIES by Glen Holstein .......................................................................... 36California’s vast prairie landscapes have been virtually ignored and so has their rich invertebrate fauna. That greatlyimpedes conservation of both.

NATIVE BEES AND FLOWERS IN CALIFORNIA PRAIRIES AND GRASSLANDS by Robbin Thorp ................ 40Remnants of once extensive California grasslands and embedded vernal pool ecosystems are home for many species ofimportant pollinators, our native bees.

RESTORING CALIFORNIA’S INLAND GRASSLANDS: DON’T FORGET THE FORBS! by Paul A. Aigner,Kristin B. Hulvey, Catherine E. Koehler, and Leslie E. Scott .......................................................................................... 41Forbs probably once dominated many of the state’s inland grasslands. Including forb species in restoration projects canenhance pollination and resistance to invasion by non-native species.

THE CNPS GRASSLAND INITIATIVE: DOCUMENTING GRASSLAND TYPES ACROSS CALIFORNIA byJennifer Buck-Diaz and Julie Evens .................................................................................................................................45CNPS’s Grassland Initiative is actively collecting and assessing data to more fully understand the diversity and variationof herbaceous vegetation across California.

CONSERVING AND MANAGING PRAIRIE, GRASSLAND, AND VERNAL POOL LANDSCAPESTHROUGH COLLABORATION by Pelayo Alvarez ..................................................................................................... 49A partnership of ranchers, environmentalists, government agencies, and researchers is working together to protectCalifornia’s grasslands.

NEW CNPS FELLOWS ......................................................................................................................... 55, 56

PETER JOSEPH CALLIZO: 1937–2011 .......................................................................................................58

BOOK REVIEWS......................................................................................................................................... 60

LETTERS TO THE EDITOR ............................................................................................................................62

THE COVER: Spectacular masses of spring wildflowers cover the slopes of the Temblor Range, Carrizo Plain. This photographcomes from Wildflower Wonders: The 50 Best Wildflower Sites in the World (Princeton University Press, 2011) and is reproducedhere courtesy of the author and photographer, Bob Gibbons.

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PRAIRIES AND GRASSLANDS: WHAT’S IN A NAME?by Glen Holstein

hen John Muir de-scended from PachecoPass into the CentralValley in 1868 and

found it a sea of flowers, he wrote,“Here it is not as in our great west-ern prairie, flowers sprinkled in thegrass, but grass in the flowers”(Worster 2008) and “All the groundwas covered, not with grass and greenleaves, but radiant corollas . . .” (Muir1992). The valley impressed Muir

not just with its wildflower abun-dance but also its grass scarcity.

In modern vegetation terminol-ogy, Muir described a valley domi-nated by forbs, not grasses. Whythen are these landscapes calledgrasslands? It is not because ofFrederic Clements, inventor of theidea they were covered with bunch-grass. He called them Pacific Prairie(Weaver & Clements 1938), but inthose days they were more fre-

quently called California Prairie(Shelford 1963) to distinguish themfrom eastern Washington’s PalousePrairie.

HOW CALIFORNIA PRAIRIEBECAME “GRASSLAND”

Grassland was not a term rou-tinely used to describe California’sprairies until Munz and Keck’s six-

Lupine (Lupinus sp.) and owl’s clover (Castilleja exserta) near Arvin, CA. Photograph from California’s Wild Gardens (University ofCalifornia Press, 2005), courtesy of photographer Craig Lippus, [email protected].

W

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page plant community classificationin their 1,681-page flora (Munz andKeck 1959) divided them into asmaller Coastal Prairie located inClements’ Coast Forest Climax anda larger inland Valley Grassland inhis Grassland Climax (Weaver andClements 1938). Ironically, CoastalPrairie is as rich in native grass asValley Grassland is poor since it hasnumerous species in genera likeAgrostis, Calamagrostis, Danthonia,Deschampsia, Elymus, Festuca, andPoa.

Munz and Keck justified theirterminology by repeating a theorystarted by Clements that originalnative bunchgrass vegetation in Cali-fornia valleys was destroyed by graz-ing and supplanted by non-nativeannual grasses. The native bunch-grass part of the theory is criticalbecause plant communities aren’tordinarily named for their weeds.For example, bluestem-sacahuistaprairie in Texas (Kuchler 1964) isn’tcalled Chinese tallowtree forest justbecause Triadica sebifera successfullyinvades it. In any case, Munz andKeck’s flora was among the very fewCalifornia botany books at that timeand consequently was the standardfor our taxonomy and, by exten-sion, our vegetation. Most Califor-nia prairies have been called ValleyGrassland or just “grassland” eversince.

IS BUNCHGRASS THEORYTRUE?

Since the “grassland” name isbased on bunchgrass theory, it’s im-portant to know if it is true. Reli-gion is faith-based but science pro-ceeds by determining if theories arefalsifiable. Muir’s observations mayseem to do that, but some wouldexplain them away. Maybe Muir justhappened to arrive in California in abrief time window between death ofthe bunchgrasses and arrival of non-native annual grasses. However, weknow that’s not true because every

early visitor to California back tothe earliest Spanish records saw justwhat Muir did. We now know thisbecause UC Riverside’s RichardMinnich compiled these extensiverecords in his 2008 book, California’sFading Wildflowers.

Thanks to Jason Hamilton(1997) and Richard Minnich we nowknow that Clements’ bunchgrasstheory was widely and quickly ac-cepted, not because of sound evi-dence, but because of his god-likestatus as an academic superstar inthe first half of the last century. Hisother theories like non-Darwinianevolution and the relatively un-changing climates needed for suc-cession to climatic climax have longbeen debunked (Hagen 1992). Onlyhis California bunchgrass theoryretains stubborn adherents eventhough we now know it lacks evi-dence.

Clements saw that parts of Cali-fornia were flat and treeless like hisNebraska home and therefore, ac-cording to his theory, were in the

same Grassland Climatic Climax.That meant they had to have thesame climate and dominant plants.No room was left for either Cali-fornia’s Mediterranean climate orMuir’s annual forbs since his theorydictated that dominants were neverannuals and were the same through-out a climatic climax. That meantsince Stipa species dominated prai-ries in Nebraska, they had to domi-nate them in California as well.Purple needlegrass (Stipa pulchra)was found to fit the bill, althoughClements initially confused it with aGreat Plains species (Hamilton1997).

We now know that today’s dis-tribution of purple needlegrass inCalifornia is the same seen by itsearliest visitors; it is rare in valleysand common in hilly blue oak(Quercus douglasii) savannas no mat-ter how much they’re grazed. Annu-als can also be dominants like Muirdescribed and as CNPS vegetationsurveys consistently find. Once againour society is recording phenomena

Common hillside daisy (Monolopia lanceolata) and common fiddleneck (Amsinckiaintermedia) in the Temblor Range, San Luis Obispo County. Photograph by Nick Jensen.

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Muir described (1992) of manymostly annual spring forbs, followedby a few in summer and then manyagain in fall.

DO NAMES MATTER?

Once Munz and Keck’s ValleyGrassland name was adopted, Muir’swildflowers became increasingly in-visible since as forbs, not grasses,they were in a plant communitywhere they didn’t belong just whenthey were increasingly crowded outby non-native plants. First thesewere grasses like wild oats (Avena

spp.), soft chess (Bromus horde-aceus), ripgut grass (B. diandrus),medusa-head (Elymus caput-medu-sae), and rye grass (Festuca perennis).But increasingly they became exoticforbs like yellow star-thistle (Cen-taurea solstitialis), winter vetch (Vi-cia villosa), and perennial pepper-weed (Lepidium latifolium). Since onsupposedly good authority all thiswas “grassland” and its only grasseswere non-native, it is no wonder itsoon got the name “non-native an-nual grassland.” Is that a problem?

By this century’s first decade ourSacramento Valley CNPS chapter

found it definitely was. First welearned that much of the floral andfaunal diversity Muir described isstill present among non-nativeswhen looked for. Second we foundthat labeling our chapter’s most ex-tensive natural landscapes non-na-tive annual grassland made them theonly ones ignored in required gen-eral plan conservation elements, andthus land development-free firezones at the sprawl bubble’s peak.To respond, we contacted local gov-ernments and pointed out theanomalous illegality of excluding somuch biologically rich land from

Purple owl’s clover (Castilleja exserta) in Carrizo Plain, San Luis Obispo County. Photograph by Nick Jensen.

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general plans and not properly nam-ing it California Prairie. Conse-quently it is no longer excluded fromthe plan conservation elements inour chapter area.

Sprawl is now mostly stoppedby problems of its own making butnot entirely. In Sacramento County’sCordova Hills, thousands of acres ofCalifornia prairie are currently atrisk because of a project ignoring itsgeneral plan conservation consider-ations but unlikely to succeed as aconsequence. Many governmentalplanners are quite receptive to re-viving the old California Prairiename for California’s open land-scapes since they’re used to newnames. So are many academics, eventhough one recently called a hill-side painted a rainbow of colors bythe same kind of flowers Muir saw“ruined landscape” because it lackedbunchgrass. Most know good sci-ence is constantly in flux. The great-est resistance is from passive sci-ence consumers with the fundamen-talist attitude, “My college textbooksaid it, I believe it, that settles it.”

DOES CALIFORNIA PRAIRIEMATTER?

It does if you care about Califor-nia native plants, since it’s the num-ber one habitat where they’ve goneextinct. It drops to third for Califor-nia Rare Plant Rank 1B species andto fourth for all lists (Skinner andPavlik 1994). This is because prairieplants tend to have wide ranges—subject to changes causing extinc-tion—while those in other habitatsare more often on lists reflectinglimited but more secure ranges.

There is great concern that CO2

release by fossil fuel burning causesdangerous climate change, so reduc-ing it in the atmosphere is a highpriority. Since plants change it toorganic carbon while growing,they’re important for slowing cli-mate change. Trees are often plantedto do this, but prairies do it better.

Temperate forests on average store8 kilograms carbon/meter squareas plant material while prairies store3, but temperate prairies store 19as soil organic matter while forestsstore only 12. Consequently aver-age prairies store 22 and temperateforests only 20. Forests also storemuch carbon above ground wherewildfires release it, whereas prairiesstore most protected underground(Schlesinger 1991).

Our prairies are also world-classscenic resources we appreciate toolittle. A Briton ranks our CarrizoPlain and Tehachapi Range as twoof the best places to see wildflowersin the world on a list weighted toplaces closer to his home (Gibbons2011). Maybe we should start giv-ing them equal respect.

REFERENCES

Gibbons, B. 2011. Wildflower Wonders:The 50 Best Wildflower Sites in theWorld. Princeton University, Prince-ton, NJ.

Hagen, J. 1992. An Entangled Bank.Rutgers University, New Brunswick,NJ.

Hamilton, J. 1997. Changing Percep-

tions of Pre-European Grasslands inCalifornia. Madrono 44(4): 311-333.

Kuchler, A.W. 1964. Potential NaturalVegetation of the Conterminous UnitedStates. American Geographical Soci-ety, New York, NY.

Minnich, R. 2008. California’s FadingWildflowers. UC Press, Berkeley, CA.

Muir, J. 1992. John Muir: The EightWilderness Discovery Books. DiademBooks, London, UK and The Moun-taineers, Seattle, WA.

Munz, P., and D. Keck. 1959. A Cali-fornia Flora. UC Press, Berkeley, CA.

Schlesinger, W. 1991. Biogeochemistry.2d ed. Academic Press, San Diego,CA.

Shelford, V. 1963. The Ecology of NorthAmerica. University of Illinois, Ur-bana, IL.

Skinner, M., and B. Pavlik. 1994. Cali-fornia Native Plant Society’s Inventoryof Rare and Endangered VascularPlants of California. 5th ed. CNPS,Sacramento, CA.

Weaver, J., and F. Clements. 1938. PlantEcology. McGraw-Hill, New York,NY.

Worster, D. 2008. A Passion for Nature:The Life of John Muir. Oxford Uni-versity, New York, NY.

Glen Holstein, 714 Lake Terrace Circle,Davis, CA 95616; [email protected]

Johnny-tuck (Triphysaria eriantha) on older alluvium at Jepson Prairie in Solano County.Photograph by Glen Holstein.

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IS THERE STILL NATIVE DIVERSITY IN CALIFORNIAGRASSLANDS?

by Megan E. Lulow and Truman P. Young

ur understanding of thenative flora of Cali-fornia’s Central Valleyis undergoing a sea

change. Although this communityhas traditionally been described asa grassland once dominated by na-tive perennial bunchgrasses, therehave been many over the decadeswho have suggested that the em-phasis on grasses was misplaced, andthat native wildflowers (non-grassherbs) were a significant componentas well (Burcham 1957; Heady 1988;Schiffman 2007; Stromberg et al.2007; John Muir 1894; Hamilton1997).

Recently these voices have risenagain, spearheaded by Glen Holsteinof the CNPS and Richard Minnich inhis book, California’s Fading Wild-flowers. Our own survey work hasrevealed a wonderfully rich flora ofwildflowers in remnant grasslandsthroughout the state, and the pagesof Fremontia are often filled withfields of magnificent wildflowers. Inaddition to their aesthetic appeal,wildflowers contribute to ecosystemfunction, including nitrogen fixation,pollinator resources, habitat struc-ture, and forage quality.

A LOST LEGACY

The Central Valley and adjacenthills of California have been soheavily invaded by non-native spe-cies, particularly annual grasses (e.g.,bromes and wild oats) and herbs(e.g., mustards and starthistle), thatfew clues remain of the original veg-etation. Hiking through most formerrangelands in the Central Valley orwithin the California Coast Range,one finds only a scattering of nativeplant species, or perhaps rare pock-ets where natives truly dominate(Noss et al. 1995). The majority ofareas, particularly those recently re-leased from grazing, have accumu-lated high levels of thatch and deca-dent growth of non-native species.

There is very little record of thecomposition of California grasslandflora prior to the invasions of non-native species, cattle ranching, andlarge-scale cultivation (but see Muirsidebar, page 9). The invasion byexotic plants has been so extensivethat formal classifications of Cali-fornia flora recognize the grasslanddominated by non-native annualsas its own distinctive subtype, oralliance (Sawyer et al. 2009). In fact,there are few plant communities onthe planet for which we have solittle confidence about their originalcomposition.

We have observed, as have oth-ers (e.g., Ayzik Solomeshch, GlenHolstein, Peter Hopkinson, pers.comm.; see also Tables 14.3 and 14.4in Bartolome et al. 2007) that inboth “remnant” sites and sites con-verted to dominance by exotic an-nual grasses, many native wildflowerspecies persist. It is clear that callingthese communities “grasslands” maybe misleading (although we will usethe term here, for form’s sake).

We have been involved in sur-veying grasslands in two regions ofCalifornia, both of which Keeler-Wolf et al. (2007) refer to as the“Valley and south coastal” type intheir classification of the state’s ma-jor grasslands. Large-scale grasslandrestorations were being planned andimplemented throughout the tworegions we surveyed: the foothills ofthe Santa Ana Mountains within theIrvine Ranch Natural Landmark inOrange County, and the Coast Rangefoothills on the western edge of theSacramento Valley in Yolo County.For both regions we sought to docu-ment the occurrence and generalabundance of upland native “grass-land” species, and to generate a ref-erence list to facilitate the enhance-ment of species diversity in grass-land restoration projects. Both re-gions consisted of sites within alarger, often degraded landscape ofgrassland, as well as smaller grass-lands that were part of an oak wood-land or scrub mosaic.

IRVINE RANCH NATURALLANDMARK

A map of native grasslands andthe extent of their degradation wasdeveloped for management planningwithin the Lomas Ridge and Lime-stone Canyon regions of the IrvineRanch Natural Landmark. NationalNatural Landmarks are designatedby the US Secretary of Interior fornatural areas in both public and pri-vate ownership. They are recognizedas outstanding examples of the natu-ral heritage of the country, along-side national parks, recreation ar-eas, and monuments.

After a fire in October 2007, ourteam of scientists visited all but the

A post-fire wildflower display representingspecies frequently found across areas withnative remnants in grassland and grasslandscrub. Photograph by Jutta Burger.

O

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least accessible grasslands in thesehills. Grasslands were defined as hav-ing less than 20% cover of woodyspecies, including both coast live oak(Quercus agrifolia) and coastal sagescrub species. At each of 475 sitesacross the hills, we developed a listof all native species, and estimatedthe percent cover of each life form(grass, wildflower, shrub, and tree).Based on floristic composition, thesesites were divided into six commu-nity types, or alliances (Figure 1).

Across all surveyed grasslands inthe Irvine Ranch Natural Landmark,there were a total of 140 native spe-cies: 99 wildflowers, 12 grasses, 1rush, 26 shrubs, and 2 trees. Mostspecies were observed at low fre-quencies, with only two species oc-curring at more than half of all sites,

and only 15 occurring at more thanone-fifth of all sites. The number ofnative wildflower species rangedfrom 5–61 across sites, and alwaysfar exceeded the number of nativegrass species (Figure 1), and alsooften exceeded the native grasses incover.

Some of the most frequent spe-cies included: fascicled tarweed (Dei-nandra fasciculata), blue dicks (Di-chelostemma capitatum), miniaturelupine (Lupinus bicolor), coastalgoldenbush (Isocoma menziesii), andcommon goldenstars (Bloomeria cro-cea). Across all surveyed grasslands,the overall abundance and speciesrichness of native wildflowers wasgreater than that of woody speciesor geophytes (wildflowers grow-ing from underground bulbs and

corms). However, the woody andgeophyte species were more com-mon and consistently found acrossmultiple grasslands.

There were substantially morenative species in ecotone grass-lands—a transition area between twoplant community types, such as na-tive grassland and woodland, or na-tive grassland and scrub. However,at Irvine Ranch Natural Landmark,ecotones contained half the cover ofthe most common native grass,purple needlegrass (Stipa pulchra),than the more open grassland. Whilethese ecotone areas are species rich,they likely serve a different functionto the ecosystem because the rela-tive abundance (or cover) of speciesis so different from that in the corearea of each plant community type.

A mosaic of grassland (background, left), scrub (foreground), and ecotone (center), plant communities common to the Irvine RanchNatural Landmark survey region. Photograph by Jutta Burger.

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dominated by blue oak (Quercusdouglasii) in this region. At theseNorthern California sites, only pres-ence or absence of native specieswas recorded. Four of the seven siteshad cattle or sheep grazing occur-ring at low levels of intensity. Ingeneral, ecotones, rocky ridge tops,and north facing slopes were betterplaces to find remnant patches thanflatter, open grassland. Nonetheless,all sites had at least 50% of theircover represented by exotic annualplant species.

Again, wildflowers dominatedthe native species lists. Across theseven sites, at least 79 native spe-

WESTERN SACRAMENTOVALLEY FOOTHILLS

We had also carried out a surveyof grassland remnants in the west-ern Sacramento Valley foothills, lim-ited to grasslands known to haveeither a native wildflower or nativegrass component to the flora, even ifcomprising only 10–15% of the to-tal plant cover at some sites (Lulowand Young 2009). Seven areas werechosen, with a total of 58 sites sur-veyed during the spring and sum-mer of 2002–2004 (Figure 2). Grass-lands were defined as having lessthan 20% cover of woody species,

cies were found in the remnantgrassland flora (10 grasses, 67 wild-flowers, 1 shrub, and 1 tree). Thenumber of native wildflower spe-cies in each area ranged from 20 to36 (Figure 2). Many of these spe-cies were widespread; 25 specieswere observed in more than halfthe sites surveyed. Geophytes ac-counted for 20% of wildflower spe-cies that occurred in at least one-half of all sites surveyed. Some ofthe more frequent wildflowers in-cluded California yarrow (Achilleamillefolium), miniature lupine(Lupinus bicolor), brodiaea (Bro-diaea spp.), valley tassels (Castilleja

Degraded grassland dominated by non-native annual grasses at Limestone Meadow in the Irvine Ranch Natural Landmark. Photographby David Olson.

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attenuata), tree clover (Trifoliumciliolatum), and hayfield tarweed(Hemizonia congesta).

NORTHERN VS. SOUTHERNSURVEYS

While the survey within theIrvine Ranch Natural Landmarkfound that most species occur atlow frequencies across a variety ofsites, the two survey areas shared 8of their 25 most common nativewildflowers (see Table 1). Whilegeophytic species represent a smallfraction of the total richness, in both

surveys they were among the morefrequent species across sites. Thisgroup of species are often not in-cluded in grassland restoration spe-cies lists and can be cryptic in rem-nant native communities, appearingaboveground primarily only afterfires or wet years. These results sug-gest this group may be an over-looked, yet significant componentof native grassland compositionacross the state.

There were conspicuous differ-ences between the two areas. In par-ticular, native clover species werefound more frequently in the west-ern foothills of the Sacramento Val-

TABLE 1. COMMON SPECIES SHARED BETWEEN SURVEY REGIONS.

Common Name Species

Fascicled tarweed, hayfield tarweed Deinandra fasciculata or Hemizonia congesta luzulifoliaMiniature lupine Lupinus bicolorBlue dicks Dichelostemma capitatumArroyo lupine Lupinus succulentusMariposa lily: splendid, Catalina, or yellow Calochortus splendens, C. catalinae, or C. luteusGum plant Grindelia camporumShort-podded lotus, strigose lotus Acmispon (formerly Lotus) brachycarpus or A. strigosus

California plantain Plantago erecta

Note: Tarweeds, mariposa lilies, and annual lotuses did not share the exact same species between regions, but had similar taxa.

Source: Lulow, et al., 2009. Post-Fire Grassland Survey on the Irvine Ranch Natural Landmark, unpublished report. Irvine RanchConservancy.

AN EARLY ACCOUNT OF THE CENTRAL VALLEY PRAIRIE

here are several accounts of the flower-rich flora of the Central Valley in the era before large-scaleagriculture, but perhaps none as eloquent as this, from John Muir:

The Great Central Plain of California, during the months of March, April, and May, was one smooth, continuousbed of honey-bloom, so marvelously rich that, in walking from one end of it to the other, a distance of more than400 miles, your foot would press about a hundred flowers at every step. Mints, gilias, nemophilas, castilleias,and innumerable compositæ were so crowded together that, had ninety-nine per cent of them been taken away,the plain would still have seemed to any but Californians extravagantly flowery. The radiant, honey-ful corollas,touching and overlapping, and rising above one another, glowed in the living light like a sunset sky—one sheetof purple and gold, with the bright Sacramento pouring through the midst of it from the north, the San Joaquinfrom the south, and their many tributaries sweeping in at right angles from the mountains, dividing the plaininto sections fringed with trees.

— from The Mountains of California by John Muir, 1894

ley, whereas more shrubs were foundin the grasslands on the Irvine RanchNatural Landmark. Greater total spe-cies richness was recorded on theIrvine Ranch Natural Landmark, butthis may be because a far greaternumber of sites were surveyed there.

GRASSLAND? FORBLAND?PRAIRIE!

In summary, throughout the sur-veyed remnant sites, the number ofnative wildflower species was fargreater than the number of nativegrass species, particularly in grass-

T

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lands with less non-native cover orecotone grasslands. Individual nativegrass species often occurred at higherabundance (cover) than wildflowers,but as a class were rarely dominant.

The persistence of native wild-flowers may offer a tool for betterunderstanding the original compo-sition of California grasslands—animportant step in restoration. Bycomparing remnant wildflower spe-cies in more highly invaded sites tothe wildflower components of grass-land communities that are more in-tact, it may be possible to recon-

struct some of their lost components,including perennial grasses.

Disturbed areas may also havesurprisingly resilient seed banks orcorm banks. In grasslands disturbedfor over a century and with virtuallyno native cover, any of the followingprocesses—clearing built up thatch,a fire, or after responsibly applyingherbicide—can stimulate native wild-flower species to germinate that werenot evident prior to the clearing (per-sonal observation, M. Lulow).

When putting together a specieslist for restoration, identifying those

species that are able to coexist withnon-native annual species can be asimportant as determining which spe-cies are most appropriate for a givensite. This is particularly true for Cali-fornia grasslands, where it is virtu-ally impossible to completely eradi-cate non-native annuals due to theextent of their invasion throughoutthe state and their prolific seed pro-duction.

Collaborative research betweenresearchers and practitioners cangreatly inform which methods andspecies are most effective for restor-ing components of California’s na-tive wildflowers to our grasslands,yet few experiments have been con-ducted. While there will be somespecies or species groups sharedamong regions, given the diversity ofspecies and physical characteristicsof the state, restoration approachesand species used should be honedfor a given region and environment.The large number of wildflower spe-cies included in these rather coarsesurveys, their functional diversity,and their aesthetic variety suggest arich palette for ecological restoration.

These surveys also remind us ofan essential question that warrantsour revisiting, namely what shouldwe call these plant communities thatare (were!) a mixture of native grassestogether with herbaceous non-grassesthat vastly outnumber the former inspecies richness, and often dominatein cover. “Grasslands” seems clearlymisleading. “Forblands” is dismiss-ive of the grasses. While some feelthat “prairie” evokes images of thetall-grass and short-grass regions ofthe Midwest, we believe that it is themost neutral and therefore most de-scriptive term, and we are increas-ingly using it to describe the originalvegetation of open habitats in theCentral Valley.

REFERENCES

Bartolome, J.W., et al. 2007. Valley grass-land. In Terrestrial Vegetation of Cali-fornia, 3rd ed., ed. M.G. Barbour, T.

FIGURE 1. SPECIES RICHNESS OF NATIVE PLANTS ATIRVINE RANCH NATURAL LANDMARK.

FIGURE 2. SPECIES RICHNESS OF WILDFLOWERS INWESTERN SACRAMENTO VALLEY FOOTHILLS.

Note: Based on floristic composition cluster and similarity analyses, six plant communitieswere identified. All sites had a minimum of 50% relative cover of non-native species.

Source: Lulow, et al., 2009. Post-Fire Grassland Survey on the Irvine Ranch NaturalLandmark, unpublished report. Irvine Ranch Conservancy.

Note: Each site had a maximum of three native grass species, and had a minimum of 50%relative cover of non-native species.

Source: Lulow and Young, 2009.

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Keeler-Wolf, and A.A. Schoenherr.University of California Press, Berke-ley, CA.

Burcham, L.T. 1957. California rangeland: An historico-ecological studyof the range resource of California.Division of Forestry, Department ofNatural Resources, Sacramento, CA.

Hamilton, J.G. 1997. Changing percep-tions of pre-European grasslands inCalifornia. Madroño 44:311–333.

Heady, H.F. 1988. Valley grassland. InTerrestrial Vegetation of California, ed.M.G. Barbour and J. Major. Califor-nia Native Plant Society, Sacramento,CA.

Keeler-Wolf, T. et al. 2007. Communityclassification and nomenclature. InCalifornia Grasslands: Ecology andManagement, ed. M.R. Stromberg, J.D.Corbin, and C.M. D’Antonio. Univer-sity of California Press, Berkeley, CA.

Lulow, M.E., and T.P. Young. 2009. Highnative wildflower richness in CentralValley “grassland” sites in the west-ern Sacramento Valley and adjacentfoothills. Grasslands 14(3):7-11.

Minnich, R.A. 2008. California’s fad-

ing wildflowers: Lost legacy and bio-logical invasions. University of Cali-fornia Press, Berkeley, CA.

Muir, J. 1894. The Mountains of Cali-fornia. The Century Company, NewYork, NY.

Noss, R.F., E.T. LaRoe III, and J.M.Scott. 1995. Endangered ecosystemsof the United States: A preliminaryassessment of loss and degradation.National Biological Service, Wash-ington, DC.

Sawyer, J.O., T. Keeler-Wolf, and J.Evens. 2009. A Manual of CaliforniaVegetation, 2d ed. California NativePlant Society Press, Sacramento, CA.

Schiffman, P.A. 2007. Species compo-sition at the time of first Europeansettlement. In California Grasslands:Ecology and Management, ed. M.R.Stromberg, J.D. Corbin, and C.M.D’Antonio. University of CaliforniaPress, Berkeley, CA.

Stromberg, M.R., J.D. Corbin, and C.M.D’Antonio, eds. 2007. CaliforniaGrasslands: Ecology and Manage-ment. University of California Press,Berkeley, CA.

Megan Lulow, Irvine Ranch Conservancy,4727 Portola Parkway, Irvine, CA 92620,[email protected]; TrumanYoung, Department of Plant Sciences,University of California, Davis, CA 95616,[email protected]

Catalina mariposa lily (Calochortus catalinae), a native grassland geophyte, found in fine soils. Geophytes were found frequently withremnant native species in both survey regions across grassland sites. Both photographs by Fred Roberts.

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UNDERSTANDING CALIFORNIA GRASSLAND ECOLOGYby Paula M. Schiffman

typical California grasslandincludes well over 100native plant species:grasses, forbs, geophytes,

perennial herbs, and subshrubs, aswell as occasional shrubs and trees.Invasive non-native species are alsopresent, particularly annual grassesthat originated in the Mediterraneanregion and became widespread soonafter European contact and settle-ment, especially oats (Avena spp.),bromes (Bromus spp.), and barleys(Hordeum spp.). Though much lessdiverse, these non-natives are thespecies that monopolize most of thespace and resources. In fact, non-natives constitute close to 100% ofthe grass cover in the majority ofCalifornia grasslands.

How is it possible that grass-lands contain about 40% of Califor-

nia’s total plant species diversity(Schiffman 2007a) and, at the sametime, are among the most thoroughlyinvaded natural ecosystems in NorthAmerica? Why haven’t the invadersdriven the natives to extinction byoutcompeting them for limited re-sources? To a large extent, thisstrange situation can be explainedby two natural factors—one extrin-sic and the other intrinsic—that ex-ert overarching control on grasslandecology: rainfall and soil disturbance.

RAINFALL

Rainfall in California grasslandsoccurs mostly during the winter andspring and varies tremendously inquantity and timing from year toyear. Some years are dry, while oth-ers are wet. In some years precipita-

tion occurs throughout the rainy sea-son, while in other years it is fin-ished in early winter or does notbegin until spring. For grasslandplants the rainfall regime is predict-ably unpredictable, and they areadapted to it.

Like native plants in other Cali-fornia ecosystems, many nativegrassland species deal with annualvariability in rainfall by producingextensive buried seed banks. Seedsproduced by generations of plantsaccumulate in the soil and may re-main dormant, hunkered down inthe seed bank for many years, wait-ing for the right set of environmen-tal triggers to induce germinationand establishment. Each grasslandspecies seems to respond to a set ofdistinctive cues, and the amount andtiming of rainfall are usually key.

A

ABOVE AND OPPOSITE TOP: Views of Carrizo Plain National Monument grasslands in different years. ABOVE: Native wildflowers includingCalifornia poppies (Eschscholzia californica), goldfields (Lasthenia californica), and owl’s clover (Castilleja exserta) are well represented.OPPOSITE TOP: Non-native annual grasses dominate, particularly hare barley (Hordeum murinum) and red brome (Bromus madritensis). Allphotographs by the author.

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A particular species’ seeds mayrespond differently to rain occur-ring during the still, warm monthsof early fall than they would to themuch colder precipitation of Janu-ary and February. As a result, thespecies composition of any givengrassland area can vary consider-ably from year to year. The combi-nations of factors responsible for fa-cilitating the emergence of a multi-tude of native species from the soilseed bank are not well understood.Nevertheless, it is very clear that theadaptations that enable native grass-land plant populations to sit tight inthe seed bank and then to grow whenclimatic conditions become optimal,also inadvertently make it possiblefor them to avoid exposure to in-tense competition for limited re-sources that non-native invaders caninflict.

This capacity of natives to pro-duce persistent seed banks also al-lows them to exploit ecological op-portunities that arise after multi-yeardroughts, which occur periodicallyin California. Unlike the natives,most invasive non-native annual

grasses do not form lasting seedbanks. Each spring their populationsare largely derived from seeds pro-duced the previous year. Duringdroughts, non-native annual grassesproduce many fewer seeds thanusual, so a few consecutive years oflow seed production can take a bigtoll on their populations. When suf-ficient rainfall does eventually re-cur, non-native grass seeds are muchless available, and consequentlythese species are unable to reoccupyso much space.

This situation provides oppor-tunities for native seeds that hadbeen biding their time in the soilseed bank during a drought, enablingthem to access considerably moreresources than usual. It is in post-drought years that grasslands oftenhave their best spring wildflowerdisplays, sometimes with vast car-pets of yellow, orange, purple, andblue. These extravaganzas of nativecolor and diversity appear to be madepossible, in large part, by the dimin-ishment of non-native annual grasspopulations. Unfortunately, this isonly a temporary condition.

Some non-native grass seeds in-variably make it through droughts,and plants from these seeds serve asthe basis for the rapid recovery ofinvasive populations. Within a yearor two they are usually once againdominant in grasslands, making ithard to imagine that there had everbeen a time when natives prevailed.Of course the native plant speciesare still present in these grasslands.Most are back underground, hidingin the newly replenished seed bank.

The great amount of year-to-yearvariability in rainfall, and the capac-ity for native species to persist outof sight in seed banks, greatly com-plicates our ability to conduct sci-entific studies in California grass-lands. Each year the composition ofthe plant community at a given sitecan be quite different, and this natu-ral variance can obscure the effectsof interactions between native andnon-native plants, relationships be-tween plants and animals, bio-geochemical dynamics of nutrientcycling, and other important eco-logical processes. This, in turn, com-plicates efforts to make science-

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matic extremes by seed banking,much as they already do. However,the long-term impact of this is un-clear, as are effects of climate changeson non-native invaders. Becausemany native species spend a greatdeal of time out of sight in seedbanks, it could be many decadesbefore it is apparent whether somespecies are positively or negativelyimpacted.

SOIL DISTURBANCE

As a phenomenon, soil distur-bance is much more predictable thanis rainfall in California grasslands,but the effects on plants are also far-reaching. Grassland terrain is char-acteristically pockmarked with dis-turbances produced by large popu-lations of small burrowing mammalssuch as pocket gophers (Thomomysbottae), ground squirrels (Spermo-philus beecheyi), voles (Microtus cali-fornicus), and kangaroo rats (Dipo-domys spp.). In fact, it has been hy-pothesized that repeated localizeddisturbances by small burrowingmammals created the terrain ofmounds and depressions that char-acterizes some grassland areas withvernal pools. Grizzly bears (Ursusarctos) were historically abundantin this ecosystem and they also pro-duced large excavations while insearch of foods such as bulbs, grubs,and ground squirrels.

The remarkable extent of soildisturbance and the profusion ofburrowing animals that produced

based management and restorationdecisions in what has become a veryrare type of ecosystem.

Nevertheless, seed banking is alife history strategy that has servednative plant species well. For morethan two centuries, hundreds ofgrassland natives have been chal-lenged by heavily invaded environ-ments and they have persisted. With-out this ability to avoid competitionwith non-natives, the native diver-sity of California’s grasslands wouldalmost certainly be far less. But whatwill the situation be in the future,given our changing climate? It seemslikely that many native species willavoid exposure to warmer tempera-tures and greater frequencies of cli-

TOP: Slope riddled with pocket gophermounds (Thomomys bottae). The grasslandbiomass was recently burned away, makingthe mounds readily visible. Disturbed soilcan help to activate seed banks of nativeplants. BOTTOM: Close-up view of pocketgopher burrow mounds with a one-meterstick.

Thousands of circular giant kangaroo rat burrow “precincts” scattered across the grasslandlandscape at Carrizo Plain National Monument. In the spring, vegetation on mounds tendsto be taller, greener, and different in composition compared to the surrounding grasslandmatrix. The line at the bottom is a dirt road.

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them were described by virtuallyevery early chronicler of California’swildlands, starting with Sir FrancisDrake in 1579. In the 1770s, Span-ish mission padres and soldiers suchas Pedro Font and Francisco Garcésrecorded similar observations, as didthe naturalist John Muir, vertebratezoologist Joseph Grinnell, botanistAlice Eastwood, and many others inthe 19th and early 20th centuries(Schiffman 2007b). Today, the land-scape-scale mosaics of disturbedpatches produced by digging ani-mals continue to be impressive, evenin the absence of grizzly bears.

Native grassland plants have co-existed with burrowing animals inthese environments for millennia.Therefore, it is not surprising thatsome natives have ruderal (distur-bance-adapted) tendencies that en-able them to exploit the churned upsoil in these natural disturbances.Certain natives have distribution pat-terns that indicate tolerances, or evenpreferences, for growing on patchesof recently disturbed soil. Examplesinclude fiddlenecks (Amsinckiaspp.), dobiepod (Tropidocarpumgracile), California lotus (Acmisponwrangelianus), as well as Californiamustard (Caulanthus lasiophyllus)and its endangered congener, theCalifornia jewelflower (C. califor-nicus).

For other native species, such asalkali sacaton (Sporobolus airoides),burrow mounds act as safe sites,where interactions with competitorsare less intense than in the surround-ing grassland matrix. The verticalmixing caused by so much animaldigging also adds dynamism to na-tive seed banks. Previously burieddormant seeds become exposedwhen new burrow mounds are cre-ated. Burrowing can move seeds inthe opposite direction as well, fromthe surface soil to greater depths.

Although the associations ofplants growing on excavated soilsfrequently include native species,these disturbed patches are typicallydominated by ruderal non-native

plants, particularly annual grasses,filaree (Erodium cicutarium), andother opportunistic species includ-ing sweet vernal grass (Anthoxan-thum odoratum) and freeway ice-plant (Carpobrotus edulis). By hop-scotching from one small disturbedsoil patch to another, prolific rud-eral invaders can rapidly occupylarge geographic areas. The chronicnature of these disturbances thenenables them to persist. It seemslikely that the vast spatial extent ofthese animal-produced disturbancesmay have facilitated the initial spreadand establishment of some invasivenon-native plants in California grass-lands. Perhaps this explains whythey were able to become ecologi-cally entrenched so quickly, beforeanyone noticed that the invasion waseven underway.

Of course the species composi-tions of California grasslands—par-ticularly ratios of native to non-na-tive plants—are also influenced bynumerous environmental factors, inaddition to variable rainfall and soildisturbance. For example, the con-sumption of plants by animals rang-ing from grasshoppers and rabbitsto antelope and elk, as well as do-mesticated livestock such as cattle

and sheep, can be significant. In ad-dition, exposure to fire—either wild-fires, which are relatively uncom-mon in grasslands because the fuelsupply is often limited, or prescribedburns done for restoration pur-poses—can also have differential ef-fects on native and non-native spe-cies. The grassland communities thatreveal themselves to us each springare a window through which we canview the responses of native andnon-native plants to this complexmix of ecological processes.

REFERENCES

Schiffman, P.M. 2007a. Species compo-sition at the time of first settlement.In California Grasslands: Ecology andManagement, ed. M.R. Stromberg,J.D. Corbin, and C.M. D’Antonio.University of California Press, Ber-keley, CA.

______. 2007b. Ecology of native ani-mals in California grasslands. InCalifornia Grasslands: Ecology andManagement, ed. M.R. Stromberg,J.D. Corbin, and C.M. D’Antonio.

Paula M. Schiffman, Department of Biol-ogy, California State University, 18111Nordhoff Street, Northridge, CA 91330-8303, [email protected]

A giant kangaroo rat (Dipodomys ingens) burrow precinct covered with disturbance adaptednon-native redstem filaree (Erodium cicutarium) and native bristly fiddleneck (Amsinckiatessellata, green foliage), center. The less disturbed intervening area is dominated bynative California goldfields (Lasthenia californica, yellow flowers).

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BETWEEN THE BLADES: THE EVOLUTION OF PLANTDIVERSITY IN CALIFORNIA’S PRAIRIES AND GRASSLANDS

by Ellen Dean

hen I walk across anecosystem dominatedby grasses, either na-tive or nonnative, my

eyes are often focused down. Thisisn’t just to avoid ground squirrelholes or gopher mounds. I am look-ing for the multitude of native spe-cies growing at my feet. While mostCalifornians see our grass-domi-nated landscapes as a green or brownblur, those of us who love plants

have trained our eyes to see the in-credible diversity of plant form andcolor hiding between the blades. Itis often the non-grass, native plantdiversity that makes one grasslandor prairie different from another.

This is especially true when ex-amining ecosystems now dominatedby non-native grasses in the interiorof California. It is the native forbsstill growing between the non-na-tive blades that give us some idea of

the botanical history of a piece ofland. Where did these interestingand beautiful native plants comefrom? Do they grow in other placesbesides California? How did theycome together to grow in this grass-land?

One of the better known scien-tific publications that sought to an-swer some of these questions isRaven and Axelrod’s 1978 publica-tion, The Origin and Relationships ofthe California Flora. This work ex-plains in relatively plain languagethat the present-day flora of Califor-nia is derived from various ancientpaleofloras that moved over the land-scape of the Americas through geo-logic time as the climate changed. Amore recent recounting of past cli-mates and vegetation by ConstanceMillar can be found in front matterof the new 2012 edition of The JepsonManual: Vascular Plants of Califor-nia. A third excellent description ofCalifornia paleobotany and themovement of species through geo-logic time can be found in SteveEdwards’ 2004 article in the journalFour Seasons.

LEFT: Bluedicks (Dichelostemma capita-tum), with its distinctive clusters of purpleflowers, grows in many types of grasslandcommunities. Dye Creek Preserve, TehamaCounty. BELOW: Flowers are about one inchin diameter. Both photographs by MargaretStarbuck.

W

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What these works tell us is thatthe land formation, latitude, and cli-mate of what became California haschanged substantially over geologictime. Fossil evidence indicates thatmany modern conifer families, in-cluding relatives of our coast red-wood (the genus Sequoia), were rec-ognizable and present in NorthAmerica by 200 million years ago.Flowering plants were present nearwhat would become California ap-proximately 100 million years ago,but their diversity was relatively lowuntil the Tertiary Period, which be-gan 65 million years ago.

Fossils reveal that early in theTertiary Period the climate in theregion of California was warmer,with year-round rainfall. The plantsthat grew in this climate, such ascycads, figs, and magnolias havebeen extinct in California for mil-lions of years. As the Tertiary Periodprogressed, the present-day land for-mation, soils, climate, and vegeta-tion of California changed, and ourpresent-day flora took shape, butthe road was full of bumps and turnsand ups and downs. Mountains werebuilt due to plate tectonics and vol-canism, and mountains eroded. Asthe land forms changed, specializedsoil types such as serpentine, gyp-sum, alkaline plains, and vernal poolhardpans were created or exposed.Some epochs were drier, somerainier, some hotter, some colder,but overall the climate became drierand cooler.

PLANT EVOLUTION ANDMIGRATION THROUGHTIME

As the climate and habitatschanged, new types of plants fromother areas with cooler or drier cli-mates migrated into California, re-placing those that could no longertolerate the climate. As the SierraNevada rose, it became a barrier tomigration from the east, creating anisolated region where the plants

evolved into new forms unique toCalifornia. The “California Current,”an ocean circulation pattern thatexists today, began to develop ap-proximately 15 million years ago.This current was the driving forcebehind the development of the sum-mer drought Mediterranean climateregime that has characterized mostof California for the past four toseven million years. This climate,

along with the Sierra Nevada bar-rier, created “a climatic island”unique in North America. Accom-panying this change in climate wasa sorting and redistribution of plantspecies. Those that remained in re-gions of California with summerdrought (and without surface wateror fog) needed to be able to with-stand six months without water.

Many of the herbs that we

Goldfields, lupines, and poppies create patterns among the volcanic rocks of Table Mountain,Butte County. Snow Mountain can be glimpsed across the Central Valley in the background.Photograph by Margaret Starbuck.

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admire in the grasslands of the inte-rior foothills and Central Valley ofCalifornia evolved an annual life his-tory that allowed them to wait outthe drought as seeds. Others becameperennial herbs that died back tounderground stems (bulbs, corms,or rhizomes). Most of these herbssprout, flower, and fruit in thespring, before the drought is at itspeak. A major exception to this pat-tern is several genera of tarweeds.These plants have developed the ca-pacity to flower and fruit at theheight of the drought before dyingback until the next year.

About 2.6 million years ago, just

when they thought things couldn’tget worse, California’s plants werehit with the Ice Age. During theglaciations (which each lasted40,000 to 100,000 years), sea leveldropped, the climate cooled, and gla-ciers formed at high elevations. Gla-cial melt eroded hillsides, filled riv-ers, and formed lakes in the CentralValley. During the interglacials (eachlasting about 10,000 years), the cli-mate warmed and the sea level rose.Although most of California wasunglaciated, the climate fluctuationsof the glacial/interglacial cycles werechallenging for plants, because theyhad to use their seeds or otherpropagules to migrate north andsouth or up and down in elevation.In addition, the landforms of Cali-fornia were continuing to changedue to sea level changes, plate tec-tonics, volcanism, and erosion.

During the Tertiary Period andthe Ice Age, plants migrated fromplace to place and occupied newlyopened habitats. Species adapted toone soil type might find themselvespreadapted to the harsh conditionsof a newly available environment.Some plants became isolated on pen-insulas, ridges, vernal pools, or spe-cialized soil types. It is well known

that isolated populations sometimesspeciate (see Grant 1981, for ex-ample). This partly explains why wehave so many genera with speciesthat evolved in and are unique toCalifornia.

ORIGINS OF CALIFORNIA’SPRAIRIE WILDFLOWERS

After that very long introduction,shall we return to the original ques-tions about the wildflowers in thegrass? Let us think about those herbsin our interior prairies with a well-developed Mediterranean climate.Where did all these plants comefrom, do they grow in other placesbesides California, and how did theycome together in these habitats?

Daniel Axelrod hypothesizedthat many of our lower-elevation,interior California native plants camefrom more arid climates to the southand southwest. This would meanthat many of the genera present inthis region would also be presentoutside of California, especially inthe arid western US or Mexico. How-ever, some of the California speciesbelonging to those genera would berecently evolved species that are en-demic to California. Let us look atsome of the most common speciesof the interior grassland and see ifwe can understand where they mayhave come from.

Shall we start with our stateflower? Our poppies (genus Esch-scholzia) include a number of spe-cies, such as frying pans (Eschs-cholzia lobbii) and foothill poppy(E. caespitosa), as well as our Cali-fornia poppy (E. californica). Pop-pies are a vibrant component ofmany California prairies and grass-lands. The closest relative of ourpoppies are the Mexican tulip pop-pies in the genus Hunnamannia.Mexican tulip poppies are found inthe Sonoran desert of Mexico. There-fore, is it likely that our poppies inthe genus Eschscholzia migrated intoCalifornia from the south.

Hayfield tarweed (Hemizonia congesta) is alate-flowering prairie species found in thehot, summer interior of California.Photograph by Margaret Starbuck.

Poppies and birds-eye gilia (Gilia tricolor) grow in profusion in the prairies at Bear CreekRanch in Colusa County. Photograph by Craig Thomsen.

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Similarly, bluedicks (Dichelo-stemma capitatum), a lily thatemerges each spring from a corm, isthought to have evolved from anancestor that also gave rise to sev-eral species of southern lilies (Piresand Sytsma 2002). Notable amongthese relatives are the beautiful,mostly Mexican genera Milla andBessera. Therefore, this is anothergood example of a genus that prob-ably originated in arid regions to thesouth. Bluedicks has two subspe-cies. One is confined to the Califor-nia Floristic Province (the region ofwestern North America from north-ern Baja to southern Oregon with aMediterranean climate). The otherspecies is more widespread in thearid western US.

Recent research on the evolu-tion of another group of perennialherbs, our brodiaea lilies (the genusBrodiaea), indicates that they shareda common ancestor with bluedicks

(Pires and Sytsma 2002). Brodiaeas,of which there are about 18 speciesthat are mostly endemic to the Cali-fornia Floristic Province, are a verytypical prairie plant. Brodiaea wascited by Axelrod as an excellent ex-ample of a genus that evolved rela-tively rapidly in geologic time inresponse to the climatic, soil, andtopographic changes that occurredin California. Once they separatedfrom their common ancestor withbluedicks, they radiated quickly intomany different species found on anumber of specialized soil types.

Another very common genusfound in many different interior prai-rie habitats is Thysanocarpus, whichcontains the annuals fringe pod, lacepod, and spoke pod. The genus hasfive species, four of which are foundin California and one that is en-demic to Baja California. Our mostcommon California species, fringepod (Thysanocarpus curvipes), is also

ABOVE AND BELOW: California has a high diversity of clovers. A typical prairie plant list often has at least five species. White-tipped clover(Trifolium variegatum), shown here, grows in many regions of California, preferring moist swales or vernal pool edges within prairiehabitat. Top photograph by Spencer Dukstra. Bottom photograph by Margaret Starbuck.

the most widely ranging species, dis-tributed throughout the western U.S,British Columbia, and Mexico. Thethree other California species are allrestricted to the California FloristicProvince. This is another good ex-ample of a California prairie genusthat appears to have had southern

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origins. The most recent data showthat the earliest diverging speciesbranch is that of the arid Baja Cali-fornia endemic, Thysanocarpuserectus (Alexander et al. 2010).

An important and rather spec-tacular group of prairie wildflowers,in terms of yellow splashes of colorand sheer numbers, is the goldfieldgenus (Lasthenia). Goldfields aremostly annual herbs (18 species)that occupy a wide range of prairiehabitats ranging from vernal poolsto dry serpentine slopes. Many ofthe species are endemic to the Cali-

fornia Floristic Province. One spe-cies is in Chile (thought to be bird-dispersed), and others are distrib-uted north to Washington state. Likeour brodiaeas, it has been longthought that the shift to a Mediter-ranean climate, coupled with ourregion’s changes in topography,soils, and available habitat throughgeologic time, may have promotedrelatively recent and explosive spe-ciation in this genus.

This idea is supported by recentresearch on the evolution of gold-fields based on DNA sequence data

(Chan et al. 2001). The origins ofthe genus are not completely clearat this time, but it is likely that thegenus diverged from relatives thatwere based to the south. The closestknown relatives are Amblyopappus(a genus with one species that growson coastal dunes and bluffs in Cali-fornia, Baja California, and westernSouth America), and Baeriopsis (an-other genus with one species that isendemic to Guadalupe Island, at thesouthwest corner of the CaliforniaFloristic Province) (Baldwin et al.2002).

The goldfields are in the daisyfamily and are part of a larger groupof genera called tribe Madieae, whichhas its center of diversity in the Cali-fornia Floristic Province and con-tains many important interior Cali-fornia prairie genera, such as blowwives (Achyrachaena mollis) andtarweeds. Tarweed genera includetidytips (Layia spp.), hayfield tar-weed (we have one species, Hemi-zonia congesta), and tarweeds/tar-plants (Madia spp. and Centromadiaspp.).

As mentioned above, some gen-era of tarweeds have specialized inflowering and fruiting at the heightof our summer drought when fewother native plants are flowering.These late flowering species includethe rosin weeds (Calycadenia spp.)and yellow-flowered tarweed (Holo-carpha virgata). Among the earliesttarweed lines is the genus Raillar-della (Baldwin 2006). Raillardellasare now found in high-mountainenvironments in the California Flo-ristic Province and Nevada. There-fore, the immediate ancestor of thetarweeds probably did not comefrom the south, but rather frommountain regions.

Then there are the legumes inthe family Fabaceae. California isrich in legume species, and it wouldbe hard to imagine our prairies andgrasslands without our lupines(Lupinus spp.) and clovers (Trifo-lium spp.). The evolution of bothgenera is very complicated, as both

Blow wives (Achyrachaena mollis) is a very common, annual prairie species, but it is rareto see it in the profusion shown here in the Cache Creek Wilderness, Lake County, whereit is growing with common tarweed (Madia gracilis) and Ithurial spear (Triteleia laxa).Photograph by Craig Thomsen.

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are present in the Old and NewWorlds, include annuals and peren-nials, and occupy a wide variety ofhabitats. The North American lu-pines form a distinct group. Inter-estingly, two of our most commoninterior California prairie plants,chick lupine (Lupinus microcarpus)and miniature lupine (Lupinus bi-color), are near the bottom of theNorth American lupine evolution-ary tree, and they are most closelyrelated to Mojave lupine (Lupinusodoratus) whose distribution in-cludes the Mojave Desert and BajaCalifornia (Eastwood et al. 2008).

The evolution of our clovers ismore complicated. The Californiaspecies are part of the North Ameri-can diversification of the genus, andour interior prairie species havemultiple origins, from the south,west, and north (Ellison et al. 2006).

To finish up our discussion ofthe origins of our interior Californiaprairie plants, let us look at one ofthe most common and widely dis-tributed species, California plantain(Plantago erecta). This wonderfullittle annual plant completes its lifecycle each spring in a matter ofweeks. It is endemic to the Califor-nia Floristic Province. Like lupineand clover, the genus Plantago hasspecies worldwide; some of the Eu-ropean species are part of the non-native flora of California (like theplantains in your lawn). Researchon the evolution of plantain indi-cates that New World plantainsevolved from Old World plantains(Ronsted et al. 2002). Californiaplantain is part of a group of speciesthat occur both in the eastern USand South America. Therefore, likeour clovers, it is somewhat difficultto understand where this importantspecies came from.

When we look at the origins ofthese beautiful plants that grow inour interior, Mediterranean climateprairies, it appears that many haveprobably evolved from arid-climateancestors that moved into Califor-nia from the south. However, re-

searchers are finding that some havemore complicated histories, includ-ing migration from the north, fromhigher elevations, or perhaps on abird’s foot. Some of our interior prai-rie species have broad distributionsthat include many other westernstates. Others are endemic to theCalifornia Floristic Province, andsome may be endemic to just a smallregion or confined to a particularhilltop of California. If we observecarefully and investigate, each spe-cies has a unique story to tell. Thewildflower assemblages growingwithin the native and non-nativegrasses of our prairies are but snap-shots in geological time.

REFERENCES

Alexander, P.J., et al. 2010. Molecularphylogenetics and taxonomy of thegenus Thysanocarpus (Brassicaceae).Systematic Botany 35:559-577.

Baldwin, B.G., B.L. Wessa, and J.Panero. 2002. Nuclear rDNA evi-dence for major lineages of helenioidHeliantheae (Compositae). System-atic Botany 27: 161-198.

Baldwin, B.G. 2006. Contrasting pat-terns and processes of evolutionarychange in the tarweed-silverswordlineage: Revisiting Clausen, Keck,and Hiesey’s findings. Annals of theMissouri Botanical Garden 93: 64-93.

Chan, R., B.G. Baldwin, and R. Ornduff.2001. Goldfields revisited: A molecu-lar phylogenetic perspective on theevolution of Lasthenia (Compositae:Heliantheae sensu lato). Interna-tional Journal of Plant Sciences 162:1347-1360.

Eastwood, R.J., et al. 2008. Diversityand evolutionary history of lupins—insights from new phylogenies. InLupins for Health and Wealth, ed. J.A.Palta and J.B. Berger. Proceedings ofthe 12th International Lupin Con-ference, Fremantle, Western Austra-lia. International Lupin Association,Canterbury, New Zealand.

Edwards, S.W. 2004. Paleobotany ofCalifornia. The Four Seasons 12(2):3-75.

Ellison, N.W., et al. 2006. Molecularphylogenetics of the clover genus(Trifolium—Leguminosae). Molecu-

lar Phylogenetics and Evolution 39:688-705.

Grant, V. 1981. Plant Speciation. Co-lumbia University Press, New York,NY.

Millar, C.I. 2012. Geologic, climatic,and vegetation history. In The JepsonManual: Vascular Plants of California,ed. B.G. Baldwin et al. University ofCalifornia Press, Berkeley, CA.

Pires, J.C., and K.J. Sytsma. 2002. Aphylogenetic evaluation of a biosys-tematics framework: Brodiaea andrelated petaloid monocots (Themi-daceae). American Journal of Botany89: 1342-1359.

Raven, P.H. and D.I. Axelrod. 1978.Origin and relationships of the Cali-fornia flora. University of CaliforniaPublications in Botany 72: 1-134.

Ronsted, N., et al. 2002. Phylogene-tic relationships within Plantago(Plantaginaceae): Evidence fromnuclear ribosomal ITS and plastidtrnL-F sequence data. Botanical Jour-nal of the Linnean Society 139: 323-338.

Still, S. 2012. Systematics and Taxo-nomic Studies of Eschscholtzieae(Papaveraceae). PhD dissertation,University of California, Davis. Avail-able from Dissertation and Theses atthe University of California (Publi-cation No. AAT 3456870).

Ellen Dean, 708 Mulberry Lane, Davis,CA 95616, [email protected]

California’s lupines are a large, diversegroup. This annual sky lupine (Lupinusnanus) is growing in a sea of goldfields(Lasthenia spp.) on Table Mountain, ButteCounty. Photograph by Margaret Starbuck.

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GEOLOGY, CLIMATE, AND CALIFORNIAPRAIRIE DISTRIBUTION

by Glen Holstein

merican vegetation ecologybegan a little over a hun-dred years ago in the Mid-west, where its main

founding fathers including HenryCowles, Frederick Clements, andWilliam Cooper started their careers.Inevitably vegetation conditions inand near the region at that timegreatly influenced theories about itthat they developed. Forests in the

region’s wetter eastern portion ex-tended continuously from Labradorto Florida, and grasslands in its drierwest were just as continuous fromSaskatchewan to Mexico. Whetherforest or grassland, these landscapeswere mainly altered by farms lessthan 100 years old.

Cowles and others developedtheir main ecology tenets in thesekinds of vegetation. One was succes-

sion, vegetation change across time,as for example when farms are aban-doned. Another was climatic climax,Clements’ theory that succession con-verges toward a regional vegetationtype largely determined by climate.

FACTS VS. THEORY

While climatic climax theoryworked fairly well in and around the

A

Solano County hills in May with open prairie still green on clay sediments, but golden amid oak woodland on volcanic rocks. Allphotographs by the author unless otherwise noted.

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Midwest, California caused it realproblems. William Cooper saw thatmost of the state had a fairly uni-form Mediterranean climate whichclimax theory said should produceone kind of vegetation. Conformingwith theory, Cooper said this waschaparral, meaning shrubs once cov-ered all the California FloristicProvince’s warmer and drier areasincluding valleys. But unfortunatelyfor his theory, its earliest visitorsnever reported shrubs in valleys.

In contrast, Clements recognizedCalifornia’s vegetation was diverse.To fit it to his climatic climax theory,he concluded our climate was alsodiverse. The vast plains of the Cen-tral Valley may have reminded himof his Nebraska home state sincehe annexed it to his Grassland Cli-matic Climax (Weaver and Clements1938) (Figure 3). To justify that, healso claimed that the Central Valleyand Nebraska had the same cli-mates, meaning Fresno’s climate wasmore like Omaha’s than Riverside’s(Hamilton 1997).

Clements also claimed Stipabunchgrasses once covered the Cen-tral Valley, but its earliest visitorsdidn’t see these either except in ad-jacent foothills where they’re stillcommon. What the valley’s earliestvisitors did see was a vast sea ofwildflowers, a phenomenon welldocumented by John Muir but alsoreported by even the earliest valleyvisitors (Minnich 2008). How thencould Clements ignore these wild-flowers? He had to because histheory had a rule that vegetationmust be dominated by perennials(Hamilton 1997). Since the valley’swildflowers were mostly annuals,Clements’ theory would not permitthem to dominate vegetation no mat-ter how many there were. Despiteits rarity in the Central Valley, hefound the ideal perennial his theoryneeded in purple needle-grass (Stipapulchra), since he initially mistookit for one of his Grassland ClimaticClimax’s abundant Great Plains spe-cies (Hamilton 1997).

CALIFORNIA’SVEGETATION MOSAIC

Climatic climax theory had a hardtime explaining California vegetation,which changed much faster acrossshort distances than climate. In anapparently common climate forest,savanna, sage scrub, chaparral, oropen prairie often coexisted in a com-

plex mosaic seen and described byCalifornia’s earliest visitors. Climaticclimax also lost support when it waslearned how fast climate changes.Nevertheless, just 17 years ago somestill interpreted the mosaic as differ-ent successional stages. Clements’claim that bunchgrass once coveredthe Central Valley—the later justi-fication for calling it “grassland”—

TOP: White layia (Layia glandulosa) on recent alluvium of Central Valley floor in KernCounty, March 2011. • BOTTOM: Ithuriel’s spear (Triteleia laxa) on uplifted older alluvium,and riparian vegetation (in background) on recent alluvium at Tassajara Hills, ContraCosta County.

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is even more enduring. Elna Bakker(1971), an observant non-academic,was the first to treat our mosaic as anatural phenomenon and at least par-tially explain its occurrence here.

Why is California different fromthe eastern United States? One rea-son is its Mediterranean climate. Eastof the Rockies it rains in summerwhen plants grow fastest, but in Cali-fornia the wet and growing seasonsare months apart, a period whenwater must be stored in soils androcks to be used when the growingseason’s long, warm days arrive. Eu-ropean vegetation ecologists likeDenmark’s Eugenius Warming in1895 gave such phenomena result-ing from geology’s effect on vegeta-tion much more attention than didour Midwestern ecologists, who hadfewer relevant local examples. Morerecently Germany’s Heinrich Walter(1979) has been particularly insight-ful in explaining how geology andclimate interact to create vegetation.

Southern France has a Mediter-ranean climate less extreme thanCalifornia’s, but vegetation that simi-larly changes rapidly across space.It was there Josias Braun-Blanquetdeveloped a method for describingvegetation that is now used world-wide, including by the CNPS Veg-etation Program, which identifiesareas with distinctive vegetationcalled relevés and then estimatestheir coverage by different species.

In summary, the US and Europedeveloped quite different emphaseson how and why vegetation changes.Americans tended to focus on veg-etation change in time through suc-cession to a climatic climax, whileEuropeans focused on change acrossspace caused by multiple environ-mental factors, including geology.

WHERE ARE CALIFORNIA’SPRAIRIES?

In 1957 range ecologist L.T.Burcham created a very good map ofCalifornia’s pristine vegetation with-

out the benefit of modern satelliteimagery (Burcham 1957) (Figure 1).He mapped as “California Prairie”areas lacking dominant woody plantswhich occur throughout California’sMediterranean climate zone in awide range of local climates.

California prairie occurs inHumboldt County where rainfallexceeds 70 inches and in the Cuyamaand southern San Joaquin Valleyswhere it averages only 5. It is inShasta Valley where the Januarymean temperature is 34º F andthe mean minimum is 24º F, and inthe Palos Verdes Peninsula whereequivalent temperatures are 57 and50. Prairie can be found on the PointReyes Peninsula where the Julymean is 54º F and the mean maxi-mum is 58º F, and in Bakersfieldwhere the equivalent temperaturesare 84 and 99. California prairiethus occurs across the Californiamediterranean zone’s full range ofclimatic diversity.

It is evident from Burcham’s mapthat his California prairie is associ-ated with environmental factorsother than climate, and comparisonwith geological maps indicates it isclosely associated with areas whereclay is abundant. That’s not surpris-ing since European vegetation ecolo-gists like Walter (1979) explain indetail why clay favors herbaceousplants such as grasses. Their rootsystems tend to be shallower anddenser than those of woody plantsso they depend more on upper soillayers. When these are rich in claythey hold more water and restrictdeeper movement of water and air.For this reason, clay soil favorsgrasses and other herbs, whilecoarser soil favors woody plants.

Examples of this effect on veg-etation are numerous. In Texas the12 million-acre Blackland Prairie onclay is surrounded on all sides bypost oak (Quercus stellata) wood-land on sediments containing muchless clay. It is as evident closer tohome in adjacent ridges with identi-cal climates in Solano County hills.

FIGURE 1. RANGELAND ECOLOGIST L.T.BURCHAM’S 1957 MAP OF CALIFORNIA’S PRISTINEVEGETATION.

Note: California Prairie is indicated in red. Graphics byJoan Kodani.

FIGURE 2. CALIFORNIA FIRE AND RESOURCEASSESSMENT PROGRAM (FRAP) MAP OF STATELAND COVER IN 2003.

Note: Herbaceous land cover is indicated in bright yellow.It is approximately equivalent to Burcham’s Californiaprairie. Graphics by Joan Kodani.

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There prairie on Eocene marine clayis found next to oak woodland onvolcanic rock. Because clay holdswater near the surface, its extensiveprairie stays green a full monthlonger than prairie patches on thevolcanics.

Nevertheless California prairie’sclose association with clay has beenessentially ignored until now, andgoes unmentioned in standard textson California vegetation. This hugegap in understanding Californiavegetation geography likely resultsfrom the residual influence ofClements’ overemphasis on climate.While the relation of geology tovegetation is well studied at ser-pentine outcrops and vernal poolsin California, it is otherwise ignoredhere more than elsewhere in theUS. In Georgia (Chafin 2007), forexample, rare plants are classifiedby natural communities defined al-most exclusively by geology, butthis has never been systematicallydone in California even though it ismore climatically and geologicallydiverse than Georgia and has manymore rare plants.

IF CLAY IS SO IMPORTANT,WHAT IS IT?

Clay is the generic name for amineral group that has specific char-acteristics and is created from sili-cate rocks as they weather when ex-posed to water. It is nearly inde-structible and can remain in the en-vironment for millions of years un-til extreme heat or pressure eventu-ally convert it to metamorphic orvolcanic rocks. Meanwhile it cansolidify into sedimentary rocks andbe uplifted into high mountains. Adistinctive feature is its breakup intotiny particles readily moved by windand streams. These are called dustin air, and mud in water. Since bothcan be moved much farther thanmost non-clay minerals, they oftenaccumulate far from where weath-ering created them. Clay created in

mountains, for example, is usuallycarried farther away to valley bot-toms by erosion and stream trans-port than other minerals. Conse-quently California’s many valleys arewhere clay accumulates. It is notcoincidental they’re also whereCalifornia’s prairie vegetation wasmost extensive

Clay’s importance to plants ariseswhen its tiny particles aggregate andpack tightly together, since they havea unique capacity to tightly holdlarge quantities of water and nutri-ents and gradually release them toplant roots. Packing is so tight how-ever that movement of water and air

to deeper roots is restricted. Conse-quently, soils with lots of clay areoften described as poorly drained.

PROBLEMS WITH SOILMAPS

A word should be said about soilssince they’re where plants grow. Soilscience deals with the interface be-tween geology and plants and hasdeveloped its own terminology. Inthe United States, however, this canmake clay hard to find since our soilmaps classify many soils in a mainsequence of orders determined more

Source: Weaver and Clements, 1938.

FIGURE 3. CLEMENTS (AND WEAVER 1938) “ANNEX” THE CENTRAL VALLEY TONEBRASKA AND THE GREAT PLAINS IN THEIR GRASSLAND CLIMATIC CLIMAX.

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by relative age than geology. Enti-sols, the youngest, are followed byslightly older Inceptisols, middleaged Alfisols, and old Ultisols. InCalifornia most prairie is on this agedetermined soil order sequence, butsome is on orders not defined byage. An example is Vertisols, an or-der defined by clay that cracks. It isalmost always vegetated by prairiebut underlies just a fraction in Cali-fornia because it’s a relatively un-common kind of soil.

Clay can be any age. Some is sofresh it is not considered soil, butmost is weathered enough to beEntisols, Inceptisols, or Alfisols.Consequently these orders tell littleabout vegetation or soil materials.Entisols are just very young soilsthat may form on clay, sand, or rock,all of which can greatly influencevegetation. Mollisols, an order de-fined by dark organic matter, is char-acteristic of prairies in the Midwest

but here is most often associatedwith chaparral and oaks that darkensoil below them with shed plant ma-terial. Our prairie soils with blackorganic matter are usually Vertisolsbecause of cracking clay.

Geologic maps of parent mate-rial under soils are much better forpredicting vegetation than soil maps.In much of the US, parent materialis rock that soil-forming processessuch as weathering soften into soil.However, in much of California par-ent material is alluvium—materialeroded from uplands and carried bystreams to low places where it accu-mulates until many feet thick. Oftenit begins as clay-rich mud derivedfrom eroding upland topsoil, buteven when thickly deposited in far-away valleys it retains characteris-tics of its original source like softtexture and fertility. Since these ex-tend far beneath surface soils, allu-vium is the basis for California agri-

cultural productivity. Because ofalluvium’s great thickness, soil ero-sion is less often a problem in Cali-fornia than salinity and draught.

CENTRAL VALLEY PRAIRIES

Burcham (1957) mapped mostof what he calls “pristine CaliforniaPrairie” in the Central Valley beforeagriculture had converted muchof it to cropland. That matches theearly valley’s descriptions by visi-tors like John Muir, who describedit as a sea of wildflowers with littlegrass. He identified numerous col-orful forbs dominating it in springand a second late summer domina-tion by forbs like virgate tarplant(Holocarpha virgata). Many of itsrelatives are also prominent in thislate prairie flora, but those in thetidytips genus Layia flower in thespring prairie. Prairie forbs Muirdidn’t discuss are summer bloom-

Virgate tarplant (Holocarpha virgata) on Great Valley sequence strata in Napa County.

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ing perennials like narrow-leavedmule’s ears (Wyethia angustifolia)and perennial lupine (Lupinus for-mosus), which survive in prairierelicts where theory would expectpurple needlegrass.

Burcham didn’t map all the val-ley as prairie, however. A large areain Kern County was desert scrubdominated by allscale saltbush (Atri-plex polycarpa) and spiny saltbush(A. spinifera). Freshwater marshesdominated by tules (Schoenoplectusacutus), bulrushes (S. californicus),and cattails (Typha latifolia) werewidespread in the lowest parts ofthe Central Valley, particularly inthe California Delta, in flood basinsto its north, and at the edges offormer Tulare Lake. Riparian for-ests followed major rivers into thevalley and occasionally, as in north-ern Tulare County, broadened intowide savannas dominated by valleyoak (Quercus lobata). Farther northin Tehama and Shasta counties adifferent kind of oak savanna—one dominated by blue oak (Q.douglasii)—extended down fromsurrounding foothills to almost com-pletely cover the valley’s narrownorthernmost end.

Nevertheless the Central Valley’smost widespread pristine vegetationwas California Prairie, and fromthere it extended in all directions.We can see what happened to it in arecent land cover map (Figure 2)produced by remote sensing andother sources (FRAP 2003) since itsherbaceous cover type is roughlyequivalent to Burcham’s CaliforniaPrairie. This map shows a few prai-rie areas not mapped by Burchamand much prairie loss to urbaniza-tion and cultivation, especially inthe Central Valley.

Both maps show that most, butnot all, California prairie landscapesare on alluvium, which favors her-baceous vegetation because of itssoft texture and often abundant clay.Historically most Central Valley prai-ries were on alluvium mapped asrecent because it was deposited dur-

ing the 10,000 years since the IceAge ended. Because its soft texture,high fertility, and low relief makerecent alluvium the easiest parentmaterial to use for cultivating cropsand constructing buildings, most ofit that was once prairie is now eitherfarmed or urbanized.

Although it was originally lesscommon, most Central Valley prai-ries are now found on older allu-vium that formed from 10,000 to 2million years ago and tends to beharder, less fertile, hillier, and lo-cated farther from the valley centerthan its younger relative. Conse-quently it is less often cultivated orurbanized and is most often usedfor cattle grazing, a land use thatpreserves many native prairie plantsby minimally disturbing soil whileremoving many competitive non-native weeds. Prairies on older al-luvium along the valley’s easternedge often have silica hardpans anda microtopography of miniaturehills and valleys that preserves rem-nant channels of streams whichonce meandered across floors ofvalleys before Sierran uplift leftthem high and dry. The hardpansand microtopography created ver-nal pools that are by far the world’sfinest.

Prairie uplands surrounding ver-nal pools often receive much lessattention, however, despite provid-ing critical nesting sites for bees pol-linating pool plants, refuge burrowsfor amphibians when pools dry, andhabitat for a rich flora and fauna alltheir own. Vernal pool prairie alsooccurs along the valley’s west edgeon uplifted older alluvium, as atJepson Prairie in Solano County, butdrainage beneath its pools is re-stricted by salty clay pans instead ofhardpans. That often causes poolwater chemistry and endemic poolspecies to be different on the valley’stwo sides.

Blue oak woodland increasinglydominates on older alluvium at thevalley’s northern end as mean an-nual rainfall increases from 19 inches

at Sacramento to 39 inches atRedding. Even there, however, prai-ries occasionally occur on both olderalluvium and on parts of the GreatValley sequence, which consists of200- to 70-million-years-old alter-nating shale and sandstone marinesediments that are turned on edge,so that ridges are formed by hardsandstone and valleys between themby softer clay-rich shale. Somesmaller prairies around the north-ern valley are famous for spectacu-lar wildflower displays caused byforb dominance. These includeColusa County’s Bear Valley on re-cent alluvium and Butte County’sTable Mountain, where a 15-mil-lion-year-old lava flow’s hard ex-posed surface functions like clay tokeep vegetation herbaceous.

The Central Valley’s recent allu-vium didn’t just cover the valley’sextensive flat clay plains. It was alsofound on natural levees and majorriver floodplains, where groundwa-ter was often shallow enough to bereached by tree roots and drainagewas better in the usually coarsersoils. These landscapes were histori-cally dominated by valley oaks thatsometimes, as at Woodland in YoloCounty, grew densely enough toform forests. Where groundwaterscarcity caused greater water stress,however, oaks were often spacedwidely in savannas that includedmuch prairie. These savanna prai-ries, the Central Valley’s greatest his-toric grasslands, were frequentlydominated by beardless wild rye(Elymus triticoides).

Salt accumulation in other Cen-tral Valley recent alluvium areas pro-duced an unusual alkaline prairieoften dominated by spikeplant(Centromadia pungens), rich in en-demic Atriplex species, and some-times containing the large bunchgrass alkali sacaton (Sporobolusairoides). Alkaline prairie is mostextensive in the wide, dry SanJoaquin Valley but also occasionallyfound in the Sacramento Valley onbasin rims, the interface between

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alluvial fans and flood basins alongmajor rivers.

THE VALLEY WASN’T ALLCLAY

While clay dominates the valley’sparent material, it also has three largelocalized areas covered by sand. Thenorthernmost in eastern ContraCosta County—before its recentthorough urbanization—was cov-ered by coast live oak (Quercus agri-folia) woodland except at its north-ern end, where a unique scrublandwas present on the Antioch Dunesjust south of the Delta’s western out-let. Farther south prairies once oc-curred on sand deposited in the SanJoaquin Valley by rivers carrying itfrom sources in rocks ground to sandby Sierran Ice Age glaciers.

The largest of these southernsand areas is Merced County’s SandyMush country, where sand accumu-lated after being carried down theMerced River from glacial erosionthat created Yosemite Valley. It isidentified as sand on geologic maps,but a similar Fresno County areacovered by sand eroded from KingsCanyon is only indicated on soilmaps. Since the Sandy Mush regionwas easy to farm and converted al-most entirely to cropland early inthe valley’s history, little is knownabout its native plant life except thatit once included the now extinctendemic Merced monardella (Mon-ardella leucocephala).

Original Sandy Mush vegetationwas likely prairie since there’s noevidence it had dominant trees orshrubs, but it may have been prairiedifferent from that in the rest of thevalley. Studying vegetation of a re-cent addition to Merced NationalWildlife Refuge at Sunrise Ranch—a 2,500-acre Sandy Mush site thatwas never plowed and has relictdunes—may reveal why such sandareas had so few woody plants de-spite being halfway between eastContra Costa’s oak woodlands and

Kern County’s Atriplex shrublands.The rarity in California of SandyMush’s lack of woody plants on sandis evident to its south in westernFresno County’s Ciervo Hills, whereislands of tall shrubs on sand sur-rounded by prairie on clay are vis-ible from miles away on Interstate 5.

Sand favors trees and shrubs be-cause it’s the opposite of clay. Airand water penetrate it freely butmoisture and nutrients are scarcenear its surface. Among grasses,these conditions favor bunchgrassessince their “bunch” or tussock ex-tends roots in all directions to har-vest scarce nutrients and water. Rhi-zomatous grasses, in contrast, dowell on clay because they can denselycover its moist fertile surface (butsome rhizomatous species are fa-vored on actively blowing sand be-cause they can quickly extend tonew, more favorable locations)(Grime 2001). It is possible purpleneedlegrass formerly dominated thevalley’s sandy areas. If Clements sawthat, he may have extrapolated it tothe whole valley.

BREACHING THEMOUNTAIN BARRIER

Cismontane and the CaliforniaFloristic Province refer to Califor-nia west of the Cascade, Sierra,Transverse, and Peninsular ranges.East of them are the transmontaneSonoran, Mojave, and Great Basindeserts. Just about all CaliforniaPrairie is west of them and cis-montane. The mountains create amostly impenetrable barrier in theHigh Sierras but are leakier to theirnorth and south. This is evidentnorth of Mount Shasta at the west-ern end of the Modoc Plateau incentral Siskiyou County, where typi-cal chaparral plants like buckbrush(Ceanothus cuneatus) intermix withtransmontane species like westernjuniper (Juniperus occidentalis) andsignificant California Prairie occurson recent alluvium in Shasta Valley.

Far to the south in Kern County’ssouthern Sierra Nevada, prairie alsooccurs in a series of alluvial valleysextending from Kern River Valley inthe north to Tehachapi Valley in thesouth. These are often connectedacross intervening ridges by prairiecorridors underlain by volcanicrocks that differ from the more com-mon Sierran granite because heavierclay soils are created when theyweather.

Some of California’s most spec-tacular remaining prairies are trulytransmontane since they occur ad-jacent to the Mojave Desert on up-lifted clay-rich older alluvium fring-ing the east base of the TehachapiMountains and the north base of theLiebre Mountains. Prairie is absent,however, on sandy recent alluviumflooring the nearby Antelope Val-ley. An outstanding pristine Califor-nia prairie outlier nearby is protectedat Fairmont Butte in the AntelopeValley California Poppy State Re-serve. These prairies are geomor-phologically in the Mojave Region,but their climate is less extreme thanthe desert since transmontaneSandberg in the west Mojave prairieregion has 13 inches mean annualprecipitation, whereas cismontaneBakersfield only gets 6.

In the mountains themselvesenough clay, organic matter, andwater often accumulate in localizedbasins to suppress forest tree growthand create open meadows. These canlook like prairies and have manyplants related to those of lower el-evations, but they differ in beingmost active, green, and colorful inJune and July just when most prai-rie plants are brown and sheddingseeds. Meadows can do this becausethey receive much of their waterfrom snowmelt and thus avoid stresscaused by a Mediterranean climate.

NORTH COASTAL PRAIRIE

Prairies along California’s NorthCoast were left behind whenClements “annexed” the Central

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Valley to Nebraska. That eventuallycaused the valley’s prairies to becalled “grassland” despite havinglittle native grass, while the NorthCoast was ironically left as Cali-fornia’s only “prairie” even thoughit had a lot. Coastal prairie north ofthe Golden Gate is most extensivein the rugged hills of HumboldtCounty, and in the low ones ofnorthern Marin and western Sonomacounties between Santa Rosa andthe coast.

Humboldt prairies are most ex-tensive south of Eureka around CapeMendocino and northeast of it atBald Hills. In both areas they’re onsteep slopes and high ridges under-lain by the Franciscan Complex, amixture of sediments scraped off theocean floor that plate tectonic forceswere pushing into a subduction zonewhile the Great Valley sequence wasforming farther east. Franciscan andGreat Valley sediments both consistof much sandstone and shale, butthe former lacks the latter’s regularlayering. Franciscan sediments werethoroughly mixed during their vio-lent formation, and vegetation de-veloping on them after they wereuplifted into hills isn’t so different.On a detailed Humboldt County veg-etation map much of the area un-derlain by Franciscan is describedsimply as an oak woodland/grass-land mosaic too complex to disag-gregate, but its larger prairie areasare identifiable by mapped nests ofbirds like Western Meadowlark(Sturnella neglecta) which use themas habitat.

The mosaic isn’t continuoussince its Bald Hills and CapeMendocino areas are separated bydense redwood (Sequoia semper-virens) forest on more recent non-Franciscan sediments of the fault-bounded Eel River Basin geologicalarea. The basin was once a valley inFranciscan but is now uplifted sohigh that the Freshwater Fault at itseastern edge creates a perfect veg-etation boundary. To its west areHeadwaters redwood forest’s rugged

hills on basin sediments, and to itseast is woodland/grassland mosaicwith many meadowlark nests onequally rugged Franciscan hills. Suchevidence of geology controlling red-wood distribution should raisedoubts about its supposed fog de-pendence.

More evidence that Franciscangeology favors prairie is provided bytheir distribution in Oregon sinceboth extend north only as far as itssouthwestern-most corner in CurryCounty and then stop. Climate isunlikely to account for this sincerainfall at Gold Beach in Oregon’sFranciscan-prairie zone is 78 inchesbut only 69 at Newport farther northin coastal conifer forest with geol-ogy like the Eel River Basin. Bothtowns have only a month with lessthan an inch of rain, but summersare two degrees warmer at GoldBeach. A climate factor likely to fa-vor coastal prairie is the stress thatsalt-laden westerly winds causestrees. Forests often extend to tide-water in eastern North America sinceits prevailing westerlies usually blowfrom land, but they cross the wide,salty Gulf of Lawrence before reach-ing coastal prairie like NorthernCalifornia’s on the west coast of Cape

Breton Island. Forests extend downto saltwater, however, on its eastcoast.

California coastal prairie’s larg-est area in northern Marin and south-ern Sonoma counties is almost en-tirely on five-million-year-old Wil-son Grove formation marine sedi-ments, except for an eastern exten-sion onto non-marine older alluviumof the Santa Rosa plain. A narrowand often hilly prairie corridor on avariety of clay-rich sediments nearthe north shores of Suisun and SanPablo Bays and Carquinez Strait his-torically connected these Marin-Sonoma coastal prairies in the Cen-tral Valley.

PRAIRIES SOUTH OF THEGOLDEN GATE

South of the San Francisco Estu-ary prairie extends nearly continu-ously through the southern CoastRange from Los Medanos Hills atthe south shore of Suisun Bay to theSanta Ynez Mountains at the westend of the Transverse Range. Far-ther south it was once extensive onthe Los Angeles and Ventura coastalplains and occasionally as far as San

California goldfields (Lasthenia californica) on older alluvium at Jepson Prairie in SolanoCounty.

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Diego County. It is widespread onrecent and older alluvium in valleysfrom Livermore in the north to SantaYnez in the south, and frequentlyconnected by prairie corridors acrossintervening mountains underlainby Great Valley and more recentclay-rich marine sediments. A wideprairie corridor at Altamont Passconnecting Livermore Valley prai-ries with those of the Central Valleyacross a ridge of marine sedimentsand older alluvium typifies manysimilar southern Coast Range link-ages with the valley.

Wildflower displays are oftenspectacular on Southern Coast Rangehill and mountain prairies like thosenorth of Livermore Valley on up-lifted older alluvium at TassajaraHills dominated by Ithuriel’s spear(Triteleia laxa) and on TemblorRange marine sediments northeastof Carrizo Plain dominated by com-mon hillside daisy (Monolopialanceolata). Similar flower displaysare frequent throughout a large rela-tively unaltered prairie region ex-tending from Salinas Valley in south-ern Monterey and northern San LuisObispo counties southeastwardacross older alluvial and marine sedi-mentary ridges to broadly connectwith both the San Joaquin Valleyand Carrizo Plain, an alluvial areapartially protected at Carrizo PlainNational Monument that has prai-ries like those of the pristine Cen-tral Valley.

Diverse prairies in a triangularwestern Santa Barbara and San LuisObispo county region include bothtypical hill and valley inland prairieand Franciscan Complex coastalprairie in northwestern San LuisObispo County resembling that inNorthern California. This regionisn’t all prairie, however. Chaparral,for example, dominates sandy, clay-poor older alluvium on Burton Mesain western Santa Barbara County.Such chaparral islands surroundedby prairie or oak woodland occurthroughout the southern CoastRange wherever clay is lacking. Good

examples are extensive granite out-crops covered by chaparral in thenorthern Gabilan Range along theMonterey-San Benito county line andthe La Panza Range in central SanLuis Obispo County.

Prairies were less extensive inSouthern California except on re-cent alluvium of the now thoroughlyurbanized Ventura and Los Angelescoastal plains, but some survive onclay-rich marine sediments at OakRidge in Ventura County and atChino Hills, where Los Angeles,Orange, and San Bernardino coun-ties meet. Other Southern Califor-nia prairie areas are at RiversideCounty’s Badlands on uplifted olderalluvial clay, and in San DiegoCounty at Lake Henshaw’s alluvialvalley and areas in its coastal hillsbetween Camp Pendleton andRancho Santa Fe where volcanicrocks weather to clay.

CALIFORNIA PRAIRIE:WELCOME HOME

Climate obviously matters sinceCalifornia has deserts in the south-east where it’s hot and dry, and for-ests in the northwest where it’s cooland wet, but California vegetationoften changes abruptly even whereclimate doesn’t. In the CaliforniaFloristic Province abrupt changefrom woodland to chaparral, sagescrub, or prairie happens most oftenwhen geology changes, not climate.That suggests the province lacks amodal vegetation type determinedby climate. Woodland, chaparral,and sage scrub are well known, butprairie is hardly known at all eventhough it is often intimately associ-ated with woodland in savannas orcomplex vegetation mosaics.

Linked for too long by the name“grassland” to hypothetical bunchgrass cover that scarcely and onlylocally existed, as well as to non-native weedy annual grasses, Cali-fornia’s prairies deserve better thantheir treatment as Other in Barbour

and Billings North American Terres-trial Vegetation’s second edition,which followed Clements’ annex-ation of the Central Valley to Ne-braska by lumping them with “other”North American grasslands ratherthan including them in its two chap-ters on “real” California vegetation.It is high time they received theirrightful place in our Province’s greattriumvirate of woodland, chaparral,and prairie vegetation.

REFERENCES

Bakker, E. 1971. An Island Called Cali-fornia. University of California Press,Berkeley, CA.

Barbour, M., and W.D. Billings, eds.2000. North American Terrestrial Veg-etation. 2d ed. Cambridge UniversityPress, Cambridge, UK.

Burcham, L.T. 1957. California Range-land. California Department of Natu-ral Resources, Sacramento, CA.

Chafin, L.G. 2007. Field Guide to theRare Plants of Georgia. State Botani-cal Garden of Georgia, Athens, GA.

FRAP. 2003. Land Cover. CaliforniaDepartment of Forestry and Fire Pro-tection, Sacramento, CA.

Grime, J. P. 2001. Plant Strategies, Veg-etation Processes, and Ecosystem Prop-erties. 2d ed. John Wiley, Chichester,UK.

Hamilton, J.G. 1997. Changing percep-tions of pre-European grasslands inCalifornia. Madroño 44(4): 311-333.

Hunter, J., et al. 2005. Atlas of the Breed-ing Birds of Humboldt County, Cali-fornia. Redwood Region AudubonSociety, Eureka, CA.

Jennings, C. 2010. Geologic Map ofCalifornia, Updated Version. Califor-nia Geological Survey, Sacramento,CA.

Minnich, R. 2008. California’s FadingWildflowers. UC Press, Berkeley, CA.

Walter, H. 1979. Vegetation of the Earthand Ecological Systems of the Geo-Bio-sphere. 2d ed. Springer-Verlag, NewYork, NY.

Weaver, J.E., and F.E. Clements. 1938.Plant Ecology. McGraw-Hill, NewYork, NY.

Glen Holstein, 714 Lake Terrace Circle,Davis, CA 95616; [email protected]

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VERTEBRATES OF CALIFORNIA GRASSLANDSby Gene R. Trapp

o what degree do California’snative vertebrates—amphib-ians, reptiles, birds, and mam-mals—depend on grasslands

for their survival? If they dependentirely on grasslands to survive,then they are obligate users. If not,they are called facultative users. Butto what extent do they need grass-lands? Here we will consider notonly the obligate species, but alsothe facultative species that needgrasslands to such a large extent thatwe could call them “almost obligate.”

AMPHIBIANS

California Tiger Salamander(Ambystoma californiense)

Almost obligate. This species(7.6–12.7 cm long) is nocturnal,spends most of the year under-ground, and usually breeds in ver-nal pools or similar ponds that dryup in summer. Some of these poolsoccur in oak savanna, others on thegrassy edges of mixed woodland andlower elevation coniferous forest.Many pools have been lost to agri-culture and development. This spe-cies has lost 55% of its historic rangeand is threatened by further agricul-tural and vineyard growth, alongwith the loss of ground squirrel andpocket gopher burrows in which itsummers. Introduced bullfrogs(Lithobates catesbeianus) are seriouspredators of eggs and young. TheCalifornia tiger salamander is listedas a Federal and California Threat-ened Species.

Western Spadefoot(Spea hammondii)

Almost obligate. Cat-like pupilscharacterize this toad (3.8–6.3 cm),together with a single black sharp“spade” on each hind foot. This isused to dig backwards into soft sub-strate to make a burrow in which it

spends the dry season. Burrows ofother animals are also used. Its dis-tribution includes the Central Val-ley, associated foothills, and theSouth Coast Ranges to the coast.This toad also spends most of theyear underground. It favors openareas of short grasses in sandy orgravelly soils found in valley andfoothill grasslands, open chaparral,and pine-oak woodlands. It breedsin quiet streams and temporaryponds. This species is extinct in al-most 80% of its traditional habitatalong the south coast of California.It has also lost extensive temporarypool habitat in the Central Valleydue to agriculture and development.The western spadefoot toad is a Cali-fornia Species of Special Concern.

There are six other species of Cali-fornia amphibians that can be calledfacultative (see sidebar, page 32).

REPTILES

Blunt-Nosed Leopard Lizard(Gambelia sila)

Almost obligate. This endemiclizard (7.5–12.5 cm) occurs in the

San Joaquin Valley region, includ-ing the Carrizo Plain. Its habitatsare semiarid grasslands, alkali flats,and washes. It is no longer presentover much of its former range dueto agriculture, development, and off-road-vehicle activity. It is classifiedas a Federal and California Endan-gered Species.

There are 15 other species of Cali-fornia reptiles that can be called fac-ultative (see sidebar, page 32).

BIRDS

Mountain Plover(Charadrius montanus)

Obligate. In spite of its name,this bird breeds in the dry short-grass region of the western GreatPlains, and winters in the dry grass-lands and semideserts of California’sCentral and Imperial Valleys, Texas,and northern Mexico. The largestconcentrations are in the ImperialValley. Its population has been de-clining for decades, likely due todisturbance by agriculture, and pos-sibly pesticides. Its diet is mainlyinvertebrates.

T

Western spadefoots (Spea hammondii) arerarely seen in California grasslands. Theycome out for a short period, after it rains,to breed, but then spend the rest of theyear in earth-filled burrows. Photographby Gary Nafis.

The population size of the blunt-nosedleopard lizard (Gambelia sila) dependsupon the number of mammal dens andburrows available in its arid habitat. Pho-tograph by Gary Nafis.

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Grasshopper Sparrow(Ammodramus savannarum)

Obligate. This secretive, big-headed, short-tailed sparrow withan insect-like song prefers grass-land with some shrubs and no treesfor breeding habitat. It occurs alongcoastal California, plus the westernedge of the Central Valley and grassyfoothills of the western Sierra Ne-vada. It is insectivorous, and pre-fers grasshoppers. Overgrazing isoften detrimental to its habitat. Thissparrow needs open areas, as it is avisual forager. It winters in Mexicoand the southeastern US, and hasbeen in decline since the beginningof the 20th century due to habitatloss. It is a California Species ofSpecial Concern.

Western Meadowlark(Sturnella neglecta)

Obligate. This species is wide-spread from the northern GreatPlains to the Pacific Ocean. Oftensinging atop fence posts, showingoff its brilliant yellow breast with adramatic black chevron, it is a closerelative of blackbirds. Like them, itoften inserts its bill into the groundor a thick mat of grass and then

The elusive mountain plover (Charadriusmontanus) has been called the “prairieghost” due to its habit of facing away froman intruder and squatting motionless,making it very difficult to see. Photographby Ed Harper.

FACULTATIVE GRASSLAND AMPHIBIANS

The following amphibians found in California are facultative grass-land species: they use grassland to varying extents, but not 100%.

Common Name Scientific Name

Black-bellied slender salamander Batrachoseps nigriventrisCalifornia slender salamander B. attenuatusCalifornia toad Bufo boreas halophilusPacific treefrog or chorus frog Pseudacris regillaNorthern red-legged frog Rana auroraCalifornia red-legged frog R. draytonii

FACULTATIVE GRASSLAND REPTILES

The following reptiles are facultative grassland species: they usegrassland to varying extents, but not 100%.

Common Name Scientific Name

Western fence lizard Sceloporus occidentalisSide-blotched lizard Uta stansburianaCoast horned lizard Phrynosoma blainvilliiGilbert’s skink Plestiodon gilbertiSouthern alligator lizard Elgaria multicarinataNorthern alligator lizard E. coeruleaSharp-tailed snake Contia tenuisWestern yellow-bellied racer Coluber constrictorCoachwhip Coluber flagellumGopher snake Pituophis cateniferCalifornia glossy snake Arizona elegans occidentalisCommon kingsnake Lampropeltis getulaLong-nosed snake Rhinocheilus leconteiCommon garter snake Thamnophis sirtalisWestern rattlesnake Crotalis oreganus (viridis)

opens the mandibles (gapes) to ex-pose seeds or invertebrates.

Lapland Longspur(Calcarius lapponicus),Chestnut-Collared Longspur(C. ornatus), andMcCown’s Longspur(Rhynchophanes mccownii)

Obligate. Longspurs are sparrow-sized winter visitors that use prai-ries, pastures, open weedy and grassyfields, grain stubbles, shores, and

any open ground. They eat seedsand invertebrates. The Lapland long-spur breeds in the Arctic tundra.Chestnut-collared and McCown’slongspurs breed in the northernGreat Plains. All are rare in Califor-nia, especially McCown’s.

Short-Eared Owl(Asio flammeus)

Almost obligate. This owl withshort ear tufts was once widespreadin California, but agriculture has

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FACULTATIVE GRASSLAND BIRDS

The following birds are facultative grassland species: they use grass-land to varying extents, but not 100%.

Common Name Scientific Name

Turkey Vulture Cathartes auraCalifornia Condor Gymnogyps californianusWhite-tailed Kite Elanus leucurusGolden Eagle Aquila chrysaetosNorthern Harrier Circus cyaneusSwainson’s Hawk Buteo swainsoniFerruginous Hawk Buteo regalisRough-legged Hawk Buteo lagopusRed-tailed Hawk Buteo jamaicensisAmerican Kestrel Falco sparveriusMerlin Falco columbariusPrairie Falcon Falco mexicanusKilldeer Charadrius vociferusLong-billed Curlew Numenius americanusMourning Dove Zenaida macrouraBarn Owl Tyto albaLesser Nighthawk Chordeiles acutipennisCommon Nighthawk Chordeiles minorSay’s Phoebe Sayornis sayaLoggerhead Shrike Lanius ludovicianusWestern Kingbird Tyrannus verticalisYellow-billed Magpie Pica nutalliAmerican Crow Corvus brachyrhynchosCommon Raven Corvus coraxBarn Swallow Hirundo rusticaNorthern Rough-winged Swallow Stelgidopteryx serripennisCliff Swallow Petrochelidon pyrrhonotaRock Wren Salpinctes obsoletusVesper Sparrow Poocetes gramineusLark Sparrow Chondestes grammacusRed-winged Blackbird Agelaius phoeniceusBrewer’s Blackbird Euphagus cyanocephalus

reduced its range dramatically, sonow it breeds in scattered areasfrom Central California northward.Brown with light spots, it is activeday and night. Its habitat is opencountry: prairie, coastal grasslands,agricultural fields, fresh and saltwa-ter marshes, and shrub thickets.Voles (Microtus) are its primaryprey, plus any other mammals orbirds that it can catch. It is a Cali-fornia Species of Special Concern.

Burrowing Owl(Athene cunicularia)

Almost obligate. Active day andnight, this small brownish owl isfound throughout much of Califor-nia except the northwestern part ofthe state and at higher elevations inthe mountains. It is a species of openterrain and short vegetation: grass-lands, steppes, and deserts. Theseowls are opportunistic feeders, andeat primarily larger insects and smallmammals, but also any other verte-brates that they can catch. This is aCalifornia Species of Special Con-cern due to loss of habitat to agri-culture. Burrowing owls need to useburrows made by other animals, es-pecially California ground squirrels(Spermophilus beecheyi), so groundsquirrel poisoning campaigns resultin fewer available burrows and thusfewer owls.

Horned Lark(Eremophila alpestris)

Almost obligate. Found year-round throughout the state exceptin the northwest corner, it needsopen country of short vegetationor bare ground: prairies, deserts,steppes, and alpine habitat. It is asparrow-sized bird with a yellow faceand throat, plus a black mask andbib, and two tiny black tufts on itscrown. Seeds and insects are its pri-

Pairs of burrowing owls (Athene cunicu-laria) need short vegetation around theirden to help them spot predators and seepotential prey. There many mouths to feedhere, with one adult second from the right.Photograph by Jo Ellen Ryan.

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mary food. It is the only true larknative to North America.

Savannah Sparrow(Passerculus sandwichensis)

Almost obligate. This small spar-row occurs year-round along the

California coast, but only in sum-mer in the northern one-third of thestate and the Sierra Cascade ranges.It is a species of open country: grassymeadows, coastal marshes, dunegrasses, sedge bogs, edges of saltmarshes, cultivated fields, and lightly

grazed pastures. It has dark streaksdorsally on a brownish background,and either a yellow over-the-eye line,or a yellow spot between the eyeand the beak.

There are 32 other species of Cali-fornia birds that can be called facul-tative (see sidebar, page 33).

MAMMALS

Nelson’s (San Joaquin)Antelope Squirrel(Ammospermophilus nelsoni)

Almost obligate. This endemicground squirrel is tan with a lateralwhite stripe, and can often be seendashing between spots of shade ona hot day with its tail bent up overits back. It is 21.8–24 cm long, andis found in grassy areas, sometimeswith scattered shrubs, in the west-ern San Joaquin Valley and adja-cent valleys, including the Carrizoand Elkhorn Plains. An omnivore,it eats vegetation, fungi, insects, andseeds. It is a California ThreatenedSpecies due to loss of habitat toagriculture, urbanization, and oildevelopment.

San Joaquin Pocket Mouse(Perognathus inornatus)

Almost obligate. This small, or-ange-brown, long-faced mouse hasfur-lined cheek pouches and a mod-erately long tail, and is a nocturnaleater of seeds and insects. It is 12.8-

FACULTATIVE GRASSLAND MAMMALS

The following mammals are facultative grassland species: they usegrasslands to varying extents, but not 100%.

Common Name Scientific Name

Ornate shrew Sorex ornatus

Broad-footed mole Scapanus latimanus

California myotis Myotis californicus

Western Small-footed myotis Myotis ciliolabrum

Yuma myotis Myotis yumanensis

Canyon bat (Western pipistrelle) Parastrellus hesperus

Big brown bat Eptesicus fuscus

Western yellow bat Lasiurus xanthinus

Western red bat Lasiurus blossevillii

Hoary bat Lasiurus cinereus

Spotted bat Euderma maculatum

Townsend’s big-eared bat Corynorhinus townsendii

Pallid bat Antrozous pallidus

Brazilian free-tailed bat Tadarida brasiliensis

Western bonneted (greater mastiff) bat Eumops perotis

Desert cottontail Sylvilagus audubonii

Black-tailed jackrabbit Lepus californicus

California ground squirrel Spermophilus beecheyi

Botta’s pocket gopher Thomomys bottae

Heermann’s kangaroo rat Dipodomys heermanni

California kangaroo rat Dipodomys californicus

Stephen’s kangaroo rat Dipodomys stephensi

San Joaquin kangaroo rat Dipodomys nitratoides

Western harvest mouse Reithrodontomys megalotis

American deer mouse Peromyscus maniculatus

California vole Microtus californicus

Coyote Canis latrans

Kit fox Vulpes macrotis

Red fox Vulpes vulpes

American badger Taxidea taxus

Striped skunk Mephitis mephitis

Elk Cervus elephus

Pronghorn Antilocapra americana

Nelson’s (San Joaquin) antelope squirrelcan get all the water it needs from digestionof its foods. It does not require free water.One of its principle predators is the kit fox(Vulpes macrotis). An adult has strongermarkings than this juvenile. Photograph byDavid Rosen.

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The best place to see the endemic giant kangaroo rat (Dipodomys ingens) is on the CarrizoPlain, in the desert grasslands of southwestern San Joaquin County. It is a Federal andCalifornia Endangered Species, as is its primary predator, the kit fox (Vulpes macrotis).Photograph by David Rosen.

16.3 cm long, and gets the water itneeds when digesting food. It is en-demic to the Central Valley fromSutter Buttes to the Carrizo Plainand the Salinas Valley. It prefers drygrassland, oak savannah, and desertscrub. One subspecies is a Califor-nia Subspecies of Special Concern,the Salinas pocket mouse.

Giant Kangaroo Rat(Dipodomys ingens)

Almost obligate. This is the larg-est kangaroo rat at 31.1–34.8 cm inlength. It has a big head, large hindlegs, a long tail, and is a seed eater,and is endemic to the open desertgrasslands and scrub of the south-west San Joaquin Valley region. It isa Federal and California EndangeredSpecies, having lost much habitat toagriculture, urbanization, and oil de-velopment. It collects seeds in itsfur-lined cheek pouches and needsno free water, which it can getthrough digestion of dry seeds.

There are 33 remaining mammalspecies in California that use grass-lands to varying extents (see sidebar,page 34).

REFERENCESBat Conservation International, www.

batcon.org.Birds of North America Online. Cornell

Laboratory of Ornithology, Ithaca,NY.

CaliforniaHerps.com.Dunn, J.L., and J. Alderfer. 2011. Na-

tional Geographic Field Guide to theBirds of North America. NationalGeographic, Washington, DC.

Holing, D. 1988. California Wildlands:A Guide to the Nature ConservancyPreserves. Chronicle Books, SanFrancisco, CA.

Jameson, E.W., Jr., and H.J. Peeters.2004. Mammals of California. Cali-fornia Natural History Guides, Uni-versity of California Press, Berkeley,CA.

Mammalian Species. The American So-ciety of Mammalogists. Allen Press,Lawrence, KS. http://allenpress.com/publications/journals/mmsp.

Reid, F.A. 2006. A Field Guide to theMammals of North America North ofMexico. The Peterson Field GuideSeries, Houghton Mifflin Company,Boston, MA.

Stebbins, R.C. 2003. Western Reptilesand Amphibians. 3d ed. PetersonField Guides. Houghton MifflinCompany, Boston, MA.

Western Bat Working Group, www.wbwg.org.

Gene R. Trapp, 2313 Isle Royale Lane,Davis, CA 95616, [email protected]

The San Joaquin kit fox (Vulpes macrotis mutica) is a facultative grassland user. It occursin the desert and desert grasslands of Southern California. Its prey is rodents, rabbits, andbirds of these arid lands. Photograph by David Rosen.

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INVERTEBRATES OF CALIFORNIA’S PRAIRIESby Glen Holstein

n 1980 I ascended a steep rockyslope in dense oak-pine woodlandand finally emerged in open prai-rie. I had reached the summit of

Fresno County’s Table Mountaineast of Millerton Lake, one of manyin California so named because ofreversed topography created whenlava rivers once flowing down Sier-ran canyons cooled to rock harderthan canyon sides. The hardenedlava remained as sinuous flat mesas

long after the canyons molding themeroded away. I was there to checkreports of vernal pools on themountain’s top for the CaliforniaNatural Diversity Database.

Vernal pools were there but whatcaught my eye in them weren’t plantsbut hundreds of the biggest, mostcolorful beetles I’d ever seen in Cali-fornia. Each was nearly an inch longand brightly marked by orange andblack. I expected such spectacularinsects would be easy to identify infield guides but was wrong. It tookUC Davis’ Bohart Museum to con-firm them as molestan blister beetles(Lytta molesta), a rare species wantedby the museum, so I made the longtrip back to collect more.

A SPECIES OF THE BEEPASTURES

I asked John Pinto, a species ex-pert, why it wasn’t on rare specieslists if so scarce, and he said it wasprobably because few cared aboutbeetles. It is now on lists but still notfederally or state listed as threat-ened or endangered. Recently Ilearned my 1980 collection is thelast confirmed record of this Cali-fornia Central Valley prairie en-demic, once widespread from Yoloto Kern counties. After the 1950s,when it was still common, it starteddisappearing.

It was a creature of the bee pas-tures John Muir described, wild-flower fields with little grass extend-ing for miles across the valley floor.It was tied as closely to those flowerfields as any tropical forest special-ist since it spent its adult life amongthem and may even have done somepollinating. Most remarkable, how-ever, was a life cycle of five larvalstages or instars it shared with itsfamily. The first, the triungulin, was

more long-legged and active thanother beetle larvae. It waited amongflowers till either a pollinating soli-tary bee carried it to its undergroundnest or it crawled there on its own,where it ate pollen and went throughfour molts into progressively moresedentary larvae until transformedinto an adult beetle by the fifth.

Muir’s bee pastures and creaturesdependent on them like the molestanbeetle are now largely gone. Al-though the beetle was last seen invernal pools, there is no evidence itwas historically confined there. Morelikely they were just places bee pas-tures survived longest when invad-ing weedy grasses crowded them offsurrounding uplands.

One might expect Californianswould notice the bee pasture eco-system’s loss, but they didn’t. To benoticed and mourned requires aname, and since flowers aren’t grass,Muir’s bee pastures vanished fromconsciousness when they started tobe called grasslands. Evidence of thisis widespread in California botany.A visiting Southern California aca-demic lecturing at UC Davis recentlysaid that a photo of massed rain-bow-colored native flowers in theTehachapi-Liebre region—a placebee pastures survive—depicted adegraded and ruined landscape, pre-sumably because it had no bunch-grass. A CNPS photo of massedLasthenia with no grass is captioned“goldfield grassland.” Whipsawedbetween a well-punctured bunch-grass myth and invading grass weeds,surviving bee pastures and creaturesdependent on them like the molestanbeetle lack value without a name oridentity. That includes the splendidpoppies and lupines on the newJepson Manual cover, more of thevegetation unnamed and thus un-seen.

This is disastrous for creatures

TOP: Red-eared blister beetle (Lytta auri-culata), a Southern California species thatresembles and is related to the molestanblister beetle (Lytta molesta). Photographby Jim Hogue. • BOTTOM: A Californiatrapdoor spider (Bothriocyrtum californi-cum) strikes at a tenebrionid beetle. Photo-graph by Marshall Hedin.

I

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like the molestan beetle that dependon bee pastures because Californialand-use agencies on good botan-ical authority dismiss them as “non-native annual grassland”—the onlylandscape unfit for required generalplan conservation elements. It is bet-ter for birds because their lobby isbig enough to get attention, but thoseof very different habitats are stilllumped as “grassland species” rang-ing from prairie generalists like west-ern meadowlark (Sturnella neglecta)through bare ground specialists likeburrowing owl (Athene cunicularia)that once followed herds of tule elk(Cervus elaphus nannodes) to thosethat once hunted lush valley oak(Quercus lobata) savanna grasslandsbut now use farmland like Swain-son’s Hawk (Buteo swainsoni).

PRAIRIE INSECTS

Because bee pastures and otherCalifornia prairies go unrecognized,their associated creatures like mole-stan beetle get virtually no attentionor protection and lack comprehen-sive studies. Nevertheless some prai-rie insects and other invertebratescan be identified.

Grasshoppers of order Orthop-tera, often the most abundant prai-rie insects, are generalist plant feed-ers which often range widely acrossNorth American prairies includingthose of California. Among its com-moner prairie grasshoppers are clear-winged grasshopper (Camnula pel-lucida), big-headed grasshopper(Eupnigodes megocephala), devastat-ing grasshopper (Melanoplus devas-tator), and valley grasshopper (Oe-daleonotus enigma). Big-headed isthe most exclusively Californian ofthese. Devastating grasshopper isnamed for its historic impact on veg-etation in mass migrations, but habi-tat loss has greatly reduced these.Unlike its many relatives, its life cycleis synchronized with California’snative annual plants. Other prairieOrthoptera are endemic grass deti-cids (Clinopleura melanopleura, C.

minuta, and Decticita brevicauda),carinated shield-back katydid (Ne-duba carinata), a widespread andcommon species complex that in-cludes several California endemics,California camel cricket (Ceuthophi-lus californianus), and the smallersand cricket (Ellipes minuta).

Small insect orders includebristletails of primitive Archaeog-

natha. Although abundant in manyhabitats including prairie, their spe-cies are poorly known and lack com-mon names. Predatory mantids inMantodea include minor groundmantid (Litaneutria minor) of Cali-fornian prairies. Thrips of Thysa-noptera are mostly tiny plant eatersthat include Doane’s thrips (Acan-thothrips cortices) and western flower

TOP LEFT: California ringlet (Coenonympha tullia), a prairie specialist on baby blue-eyes(Nemophila menziesii). • TOP RIGHT: Small checkered skipper (Pyrgus sriptura), an alkaliprairie specialist on alkali-mallow (Malvella leprosa). • BOTTOM: The monarch butterfly(Danaus plexippus), a milkweed specialist on narrow-leaf milkweed (Asclepias fascicularis).All photographs by Greg Kareofelas.

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thrips (Frankliniella occidentalis)that respectively eat prairie grassesand flowers, but black hunter (Lep-tothrips mali) atypically is a preda-tor of tiny insects and mites. Neur-optera includes larger prairie preda-tors like western mantispid (Clima-ciella brunnea) and many antlionspecies that wait in the conical pitsthey construct to devour trappedprey. Prairie predators in the smallorder Mecoptera include greenstigma (Bittacus chlorostigma) andwingless scorpionfly (Apterobittacusapterus), remarkable among relativesin lacking wings and flight.

Cicadas famous for loud singingare among many diverse groups inHemiptera. California prairie spe-cies include large grass cicada(Okanagana consobrina), small grasscicada (O. minuta), and red-wingedgrass cicada (Tibicinoides cupreo-sparsus). Other Hemiptera are plantphloem-feedersthat entered aminiaturizationpath leading toaphids and theirrelatives and fi-nally to scales,the only sessileinsects. Californiaprairie species inthese groups arelupine aphis (Ma-crosiphum albi-frons), Smith’sgrass mealy bug( T r i o n y m u s

smithii), fescue grass mealy bug (T.festucae), bunchgrass scale (Aclerdacalifornica), and grass root scale(Odonaspis graminis). Among larger,more active California prairie Hemi-ptera are true bugs like the preda-tory western corsair (Rasahus thora-cicus) and plant feeders like Califor-nia plant bug (Irbisia californica),Marin grass bug (I. mollipes), cen-tral California grass bug (I. setosa),black grass bug (I. solani), milkweedbug (Lygaeus truculentus), Califor-nia false chinch bug (Nysius califor-nicus), false chinch bug (N. rapha-nus), and large milkweed bug (Onco-peltus fasciatus).

The beetles of Coleoptera, the

largest insect order, include bothpredators and specialized plant feed-ers. It differs from Hemiptera in hav-ing chewing, not sucking, mouthparts and complete metamorphosisin which larvae and adults differgreatly. Other orders discussed herewith complete metamorphosis areNeuroptera, Mecoptera, Diptera,Hymenoptera, and Lepidoptera.California prairie beetles includepink glowworm (Microphotus an-gustus), a western relative of easternfireflies; molestan beetle and rela-tives like green blister beetle (Lyttachloris), small black blister beetle(L. nigripilis), and infernal blisterbeetle (L. stygica); pasture pill beetle(Amphicyrta dentipes); dimorphicflower longhorn (Anastrangalia lae-tifica); variable tiger beetle (Cicin-dela terricola); ant scarabs (Crema-stocheilus spp.); robust green chafer(Dichelonyx robusta); punctuate

blister beetle (Epi-cauta puncticollis);little bear (Paraco-talpa ursina); andlupine seed wee-vils (Tychius spp.).The Ohlone tigerbeetle (Cicendelaohlone) is a feder-ally listed endan-gered species con-fined to remnantcoastal prairiein Santa CruzCounty.

Flies of Dip-

A prairie meloid beetle (Lytta stygica).Photograph by Jim Hogue.

A prairie grasshopper (Melanoplus spp.).Photograph by Jim Hogue.

A large prairie centipede (Scolopendra poly-morpha). Photograph by Charles Hogue.

Coronis fritillary (Speyeria coronis), a violet specialist. Photograph by Jim Hogue.

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tera are another large order. An-cestrally aquatic, they increasinglyadapted to organic waste as well aspredation and parasitism of plantsand animals for larval feeding anddevelopment, but one early branchshifted to California prairie soils.These include range crane fly (Tipulasimplex) and flightless range cranefly (T. quaylii). Other Californiaprairie flies are greater bee fly (Bom-bylius major), net-veined bee fly(Conophthorus fenestratus), valleyblack gnat (Leptoconops carteri), andwild rye midge (Rhabdophaga elymi).Parasitic larvae of genera likeAphoebantus and Sarcophaga helpcontrol prairie grasshopper popu-lations.

Even more diverse among largeorders is Hymenoptera. Its primi-tive species are large active plantfeeders, but it has evolved to minia-ture gall makers and seed and eggparasites, flying carnivory, sociality,and pollen feeding, the main sourceof flower pollination. Prairie Hy-menoptera include sheath worm(Harmolita spp.) grass feeders; vel-vet ants (Dasymutilla spp.), whichare non-social flightless wasps; king-fisher wasps (Trielis alcione), un-derground nesting prairie flowervisitors; and trapdoor spider wasps(Psorthaspis planata), hunters ofCalifornia prairie trapdoor spiders.

Ants are abundant, usually flight-less social Hymenoptera in vir-tually all habitats including Cali-fornia prairie, where common antsinclude California red ant (Formicasubpolita), black harvester ant(Messor andrei), small harvester ant(Pheidole californica), and Califor-nia harvester ant (Pogonomyrmexcalifornicus). Harvester ants carryprairie plant seeds to nests beneathconspicuous ant hills. Social Hy-menoptera also include some waspsand bees. A few parasitize others,but most bees collect pollen. Not allare social, however, since solitaryground nesters are especially impor-tant for pollinating our native prai-rie plants.

The butterflies and moths of or-der Lepidoptera are almost all plantfeeding specialists. Prairie butter-flies include calippe and zerene frit-illary (Speyeria zerene and S. calippe)that use Johnny-jump-up prairieviolets (Viola pedunculata) and havethree federally listed as endangeredCalifornia subspecies. The monarch(Danaus plexippus) uses milkweeds(Asclepias spp.); and California ring-let (Coenonympha tullia), ox-eyesatyr (Cercyonis pegala), and wood-land satyr (Cercyonis sthenele) usenative prairie grasses, which are alsoused by several skipper butterflieslike common branded skipper(Hesperia colorado), Lindsey’s skip-per (H. lindseyi), sandhill skipper(Polites sabuleti), dogstar skipper(P. sonora), and woodland skipper(Ochlodes sylvanoides). Small check-ered skipper (Pyrgus scriptura) isan alkali prairie specialist. Mothshave many more prairie species, butfew have common names. Excep-tions are Clark’s sphinx (Proser-pinus clarkiae) and federally-listedas threatened Kern primrose sphinx(Euproserpinus euterpe). The gen-era Adela, Annaphila, and Schiniain particular have many mothsassociated with California prairieflowers.

OTHER PRAIRIEINVERTEBRATES

California’s prairies have manynon-insect invertebrates from ver-nal pool endemic crustaceans sur-viving only where protected fromfish by surrounding prairie uplands,to relatively harmless stripe-tailedscorpions (Paruroctonus silvestrii),centipedes (Scolopendra polymorpha)longer than fence lizards (Scelopo-rus occidentalis), and large myga-lomorph spiders like tarantulas(Aphonopelmus eutylenum and A.reversum), trapdoor spiders (Bothrio-cyrtum californicum and Actinoxiaversicolor), and sheet-web mygalo-morphs (Calisoga theveneti and C.

longitarsis) that emerge to hunt fromunderground homes. These, likeground-nesting bee pollinators, onlysurvive where prairie soils remainintact.

A FRONTIER FOR SCIENCEAND CONSERVATION

Internet searches now test thelimits of knowledge but bring upalmost nothing on California prai-rie/grassland insects/invertebrates.Insect ecosystems in Amazonian treecanopies are apparently examinedmore closely than those in our leastknown wild gardens. This may bethe first article ever published onCalifornia prairie invertebrates andis written by a botanist, not a zoolo-gist. Hopefully its undoubted im-perfections will inspire others to dobetter.

REFERENCES

Essig, E.O. 1958. Insects and Mites ofWestern North America. MacmillanCompany, New York, NY.

Hardwicke, K., and Heydon, S. 2009.Molestan Blister Beetle. Draft reportfor Yolo Natural Heritage Program,Woodland, CA.

Hogue, C.L. 1993. Insects of the LosAngeles Basin. 2d ed. Natural HistoryMuseum of Los Angeles County, LosAngeles, CA.

Powell, J., and C.L. Hogue. 1979. Cali-fornia Insects. University of Califor-nia Press, Berkeley, CA.

Powell, J., and P. Opler. 2009. Moths ofWestern North America. University ofCalifornia Press, Berkeley, CA.

Shanks, S. No date. Lytta molesta. Un-published report for California De-partment of Fish and Game NaturalDiversity Database.

Shapiro, A., and T. Manolis. 2007. FieldGuide to Butterflies of the San Fran-cisco Bay and Sacramento Valley Re-gions. University of California Press,Berkeley, CA.

Glen Holstein, 714 Lake Terrace Circle,Davis, CA 95616, [email protected]

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NATIVE BEES AND FLOWERS INCALIFORNIA PRAIRIES AND GRASSLANDS

by Robbin Thorp

LEFT TO RIGHT: Female mining bee (Andrena blennospermatis) that specializes in collecting pollen from yellow carpet (Blennosperma nanum),a California vernal pool endemic. Photograph by Dennis L. Briggs. • Worker bumble bee (Bombus vosnesenskii) foraging for pollen onCalifornia poppy (Eschscholzia californica). Generalist bees seek out a wide range of flower types. Photograph by Gary W. Zamzow.• Female sweat bee (Halictus ligatus) with its distinctive white bands, gathering pollen from gumplant (Grindelia camporum). Photo-graph by Kathy K. Garvey. • Male cuckoo bee (Triepeolus sp.) foraging for nectar on gumplant. Cuckoo bees do not gather pollen. Photo-graph by Kathy K. Garvey. • Soft-winged flower beetle (Listrus sp.), a pollen feeder, on California poppy (Eschscholzia californica).Photograph by Kathy K. Garvey.

“To make a prairie it takes a cloverand one bee,

One clover, and a bee.And revery.The revery alone will do,If bees are few.”

—Emily Dickinson (1924)

ne would hope to seemore than one cloverand one bee, and inmany places one still

may. We may never again see asmany flowers and bees as John Muir(1894) eloquently describes in Chap-ter 16, “The Bee-Pastures,” in hisbook The Mountains of California:

When California was wild,it was one sweet bee-gardenthroughout its entire length, . . .The Great Central Plain of Cali-fornia, during the months ofMarch, April, and May, was onesmooth, continuous bed ofhoney-bloom, so marvelouslyrich that in walking from oneend of it to the other, a distanceof more than 400 miles, yourfoot would press about a hun-dred flowers at every step.

Embedded in many grasslandsof the Central Valley are special eco-systems, vernal pools, renowned fortheir flower displays. The showypatches of flowers also provide foodfor many native bees whose females

specialize in collecting pollen fortheir offspring from particular ver-nal pool flowers, such as the miningbee (Andrena blennospermatis) thatcollects pollen from yellow carpet(Blennosperma nanum). Grasslandssurrounding the pools are nestinghabitats for mining bees. These beesprovide an important ecological linkwith their pollination services be-tween the uplands and the poolflora.

Goldfields (Lasthenia spp.) in-clude early spring flowering speciesrestricted to vernal pool ecosystemsand some that occur in drier grass-land areas. Most require insects totransfer their pollen. The diversityof visitors to most goldfield flowersincludes specialist bees whose fe-males collect pollen from one or afew closely related plants, generalistbees, and a variety of flies andbeetles. Often several species of spe-cialist mining bees may be foundvisiting the same goldfields at onelocation. However, not all flowervisitors are effective pollinators, in-cluding some specialist bees.

Our state flower, the Californiapoppy (Eschscholzia californica) pro-vides no nectar, but only pollen as areward to bees. Male bees rarely visitthese flowers, except to find femalesor to spend the night within theclosed corolla. Generalist bumblebees (Bombus spp.) and sweat bees

(Halictus spp.) are the main visitors,along with small pollen feedingbeetles.

In late spring, gum plant (Grind-elia spp.) attracts a diverse guild ofnative bees, mostly generalists, in-cluding male and female cuckoo bees.These bees have little or no hair, ortheir hair is scale-like and flattenedon the body surface, so they are notwell adapted to accumulating pol-len. Females also lack any pollentransport structures, so they dependon pollen collecting host bees to pro-vide a home and food for their young.

Let us hope and strive for pro-tections and restorations of our prai-ries and grasslands so that we willhave more than just one flower andone bee for our revery.

REFERENCES

Dickinson, E. 1924. The Complete Po-ems of Emily Dickinson. Little, Brown,Boston, MA. www.bartleby.com/113/(accessed March 28, 2012).

Muir, J. 1894. The Mountains of Cali-fornia. The Century Company, NewYork, NY. http://www.yosemite.ca.us/john_muir_writings/the_mountains_of_california/ (accessedMarch 28, 2012).

Robbin W. Thorp, Department of Ento-mology, One Shields Avenue, Universityof California, Davis, CA 95616, [email protected]

O

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RESTORING CALIFORNIA’S INLAND GRASSLANDS:DON’T FORGET THE FORBS!

by Paul A. Aigner, Kristin B. Hulvey, Catherine E. Koehler, and Leslie E. Scott

alifornia’s inland grass-lands range from the foot-hills of the Sierra Nevadato the interior of the coast

ranges, and extend almost all theway to the coast south of Santa Bar-bara. Also known as “valley grass-land” or “valley/south coastal grass-land” (Heady 1977; Keeler-Wolf etal. 2007), this herbaceous vegeta-tion type is defined by its currentdominance by non-native annualgrasses and forbs. (By “forbs” werefer to all herbaceous taxa not in thePoaceae, the plant family that gener-ally includes most species we thinkof as “grasses” and that includes ce-real grains, but not rushes or sedges).

PREHISTORY

Examples of California’s inlandgrasslands as they existed beforeEuropean settlement reside almostexclusively in the collective imagi-nation of native plant enthusiasts.No other vegetation type in Cali-fornia was so completely trans-formed before it could be stud-ied by modern botanists, whohave struggled to develop an ar-chetype of this almost mythicalplant community. While the ear-liest scientific speculation aboutthe composition of grasslandsbefore European arrival empha-sized the dominance of nativeperennial bunchgrasses, scien-tific consensus has recentlyshifted to the view that annualand perennial forbs were domi-nant, with perennial grassesinterspersed or occurring inpatchy dense stands (Schiffman2007).

Evidence for this view comesfrom the historical record, mea-ger as it is, and from relict na-

tive-dominated grasslands, whichare mostly restricted to unusual soiltypes across the state (Minnich 2008;Schiffman 2007). If there is one pointof certainty, it is that prehistoricgrasslands bore little resemblanceto the golden hills of annual grassesnative to Europe and Asia that domi-nate California’s inland foothills andvalleys today.

RESTORATION GOALS ANDLIMITS

Restoration of grasslands to somesemblance of their prehistoric state(preceding the arrival of Europeans)has been an expanding practice inCalifornia over the past two decades(Stromberg et al. 2007). The uncer-tainty surrounding the compositionand structure of prehistoric grass-lands guarantees that this endeavorwill always be part art and part sci-ence, but even if a detailed descrip-

tion of these grasslands did existand restoration ecologists had thenecessary resources, meticulouslyreconstructing prehistoric grasslandson anything larger than a demon-stration scale probably would havea high risk of failure.

For the reconstructed ecosystemto be self-sustaining (i.e., persistentwithout continual management in-tervention), land managers wouldneed to reverse several centuries ofenvironmental change including cli-mate warming, increased atmo-spheric nitrogen deposition, the in-troduction of non-native speciesfrom around the world, the declineof prescribed fire as a managementtool used by indigenous people, andthe decline of animals such as griz-zly bears, pronghorn, and elk thatmay have influenced grasslandsthrough soil disturbance or grazing.

In practice, land managers typi-cally define much more limited out-comes that will hopefully persist

under current and anticipatedconditions. The outcomes speci-fied for a given project are basedon best management practicesas they are understood at thetime, but also on the collectivevalues and imaginations of thestakeholders involved. Com-monly specified outcomes forrestoration projects almost al-ways include increasing theabundance of native species.However, they might also ad-dress grassland aesthetic appeal,provision of habitat for rare orendangered animal species, sup-pression of noxious weeds andenhancement of native plant di-versity, community resistance tofuture invasion by non-nativespecies, and provision of par-ticular ecosystem services such

Prior to the influx of Eurasian annual grasses, Cali-fornia’s inland grasslands were likely dominated bynative forbs as illustrated by this display of royallarkspur (Delphinium variegatum), California gold-fields (Lasthenia californica), bird’s eye gilia (Giliatricolor), slender cottonweed (Micropus californicus),and Indian clover (Trifolium albopurpureum) in apristine serpentine grassland. Photograph byCatherine E. Koehler.

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as water uptake, soil retention, pol-lination service, and carbon stor-age.

WHY INCLUDE FORBS?

Grassland restoration projectsthat involve active planting of na-tive species have usually emphasizedthe establishment of perennialbunchgrasses over annual and pe-rennial forbs (Stromberg et al. 2007).This bias is probably due to a num-ber of reasons. One is the histori-cally prevailing view that bunch-grasses were dominant in prehis-toric grasslands. Another is the factthat a small complement of bunch-grass species is common through-out the state, which makes selectionof species for a project manageableand commercial production of seedfeasible. By contrast, there exist agreater number of grassland forbs,and many have restricted ranges,which complicates their use in res-toration work. Also, bunchgrasspopulations are persistent once es-tablished, in contrast to annual forbswhose populations often fluctuatedramatically from year to year.

Still, many reasons exist to in-clude forbs in grassland restorationprojects, not the least of which isthat forbs were likely a dominantcomponent of many prehistoricgrasslands. But even if accurate re-construction of a prehistoric grass-land assemblage is not a priority,including native forbs in grasslandrestoration projects can enhancemany of the desired outcomes com-mon to projects.

At the Donald and SylviaMcLaughlin Reserve, in the innerNorth Coast Range at the junctionof Lake, Napa, and Yolo Counties,we have worked on numerous grass-land restoration projects in diversesettings. We have found that increas-ing the cover and diversity of nativeforbs—either by actively plantingthem or by removing the invasivespecies with which they compete—has advanced our restoration goals.As one example, hayfield tarplant(Hemizonia congesta) was added to anative bunchgrass seed mix specifi-cally to armor a restored grasslandagainst reinvasion by yellow starthistle (Centaurea solstitialis, here-after abbreviated “YST”). As a sec-

ond example, control of the inva-sive annual barbed goatgrass (Aegi-lops triuncialis, hereafter abbreviated“BGG”) resulted in a natural resur-gence of native forbs and a corre-sponding increase in the activity ofnative bees.

STARTING CONDITIONDETERMINES GOALS ANDAPPROACHES

The McLaughlin Reserve is amosaic of soil types derived fromserpentine (which we define broadlyto include all ultramafic rocks highin iron and magnesium), volcanic,and sedimentary parent material.Serpentine soils are not tolerated bya broad range of non-native species(e.g., YST) that have invaded grass-lands across the state. As a conse-quence, they support some of thebest relict native-dominated grass-lands in the region. Despite this,serpentine grasslands have not beenimmune to environmental change.Over the past three decades, BGGand Italian ryegrass (Festuca peren-nis) have spread rapidly into ser-pentine grasslands, and this processseems to be accelerated by nitrogendeposition from air pollution (Weiss1999).

Prior to intensive managementefforts that began at the reserve in2008, these two serpentine tolerantinvaders had formed dense stands inmany of the reserve’s serpentinegrasslands, although fortunatelythese grasslands still retained a highcomplement of native grasses andforbs. A typical serpentine grasslandat the reserve includes more than 50native forb species and 4 native grassspecies, all occurring within abouthalf an acre. By contrast, grasslandson non-serpentine soils contain fewto no native species, and are domi-nated by non-native annual grassesand forbs, particularly Italian rye-grass, wild oats (Avena spp.), bromes(Bromus spp.), medusa head (Elymuscaput-medusae), YST, black mustard

ADVANTAGE OF USING FORBS IN RESTORATION

grassland restoration project on Homestake Mining Company’s tailingsimpoundment at the McLaughlin Reserve. This project is typical of

many grassland restoration projects in California in that so far it includesonly grasses—in this case blue wildrye (Elymus glaucus), slender wheatgrass(Elymus trachycaulus), beardless wild rye (Elymus triticoides), and meadow

barley (Hordeum brachyan-therum). Invasion by yellowstar thistle (Centaurea solstiti-alis) and short podded mus-tard (Hirschfeldia incana) hasbeen managed with broad-leafherbicide applications, butadding native forbs such ashayfield tarplant (Hemzoniacongesta) could provide resis-tance to weed invasion with-out herbicides. Photograph byPaul A. Aigner.

A

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(Brassica nigra), vetch (Vicia spp.),and filaree (Erodium spp.).

We formulate goals for restora-tion in serpentine versus non-ser-pentine grasslands based on the dif-ference in their condition. Givenonly limited damage in serpentinegrasslands, preserving and restoringthe apparently prehistoric conditionis not only achievable, but exigent.By contrast, setting goals for non-serpentine grasslands is inevitably amore complex task. Because non-serpentine grasslands usually lackany significant native seed bank, re-storing them almost always requiresthe reseeding of native species. Incases where residual native speciescover is present, we can work topreserve this, managing invasive spe-cies and seeding additional natives.

More often than not, we find vir-tually no residual native plant com-munity and must adopt a “blankslate” approach. This involves oblit-erating the existing community tothe greatest extent possible (includ-ing the seed bank) and installing anative-dominated community fromscratch. Much of our grassland res-toration has occurred in partnershipwith Homestake Mining Companyof California, which mined more than3.5 million ounces of gold from thesite between 1985 and 2002 and re-moved more than 1,000 acres of na-tive vegetation in the process. Areasreclaimed after mining necessarilyrequire the blank slate approach.

FORBS REDUCE INVASION

Our restoration work withHomestake initially took a tradi-tional approach that emphasized theestablishment of native bunch-grasses. Following the earthmovingthat is employed to return soil to adisturbed site or to cover mine spoils,we applied a seed mix of up to sevennative grass species. Invasion by YSThas been the greatest impediment tosuccessful grass establishment, andannual applications of a broad leafherbicide have been necessary to

keep YST at bay. While herbicidesare effective at controlling YST, anecosystem that depends on regularherbicide application does not meetour definition of self-sustaining.

A promising technique for pro-viding natural invasion resistance torestored grasslands is to select na-tive species for planting that com-pete strongly for the same resourcesas likely invaders (Young et al.2009). Native tarplants are amongthe grassland species most similarto YST in the timing of growth andseed set. These late-season annualforbs germinate with the first fallrains and spend the winter andspring developing a tap root. Doingso allows them to take advantage ofdeep soil moisture in the summerand to flower and set seed long afterthe majority of annuals have com-pleted their life cycle.

Starting in 2008, we testedwhether including the hayfieldtarplant in a seed mix with nativeperennial grasses increased the re-sistance of the resulting communityto YST invasion compared to seed-ing grasses alone. Over three years,tarplant abundance remained highin plots where it was seeded, andYST abundance was 18% to 63%lower compared to plots with nativegrass alone. Adding hayfield tarplantto the restoration seed mix also re-duced the abundance of other na-tive, late season forbs such as yellow

USING FORBS TO COMPETE WITH INVASIVE PLANTS

ayfield tarplant (Hemzonia con-gesta) has a similar ecological

niche to the invasive yellow star thistle.Both germinate early and spend thewinter and spring developing a deeptaproot so they can access deep soilmoisture necessary for flowering andsetting fruit in the summer. Whenincluded in grassland restorationprojects, hayfield tarplant competeswith yellow star thistle and reduces its ability to establish. Photograph byCatherine E. Koehler.

H

tarplant (Holocarpha virgata) anddove weed (Croton setigerus) thatnaturally colonized the experimen-tal plots. This suggests that YST,tarplants, and other late season forbsform a guild of strongly interactingspecies. It also suggests that wemight more effectively exclude YSTif we plant a diversity of native lateseason forbs, each of which com-petes for resources with YST inslightly different ways.

FORBS INCREASEPOLLINATORS

Competition for essential plantresources such as nutrients, water,and light seems to play less of a rolein serpentine grasslands comparedto their non-serpentine counterparts.Preliminary results from our researchsuggest that the non-native invasiveBGG is a poor competitor, but isextremely tolerant of harsh condi-tions such as those found on serpen-tine. In grasslands with productivenon-serpentine soils, BGG probablyrequires a disturbance that reducescompetition in order to invade, butin low-productivity serpentine soilsBGG can readily invade an undis-turbed native-dominated grassland.We are therefore unlikely to preventBGG from invading serpentine grass-lands simply by seeding functionallysimilar native species.

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Nevertheless we have been suc-cessful at selectively eradicating BGGand Italian ryegrass from serpentinegrasslands by using a combinationof prescribed fire, grass-specific her-bicides, and mowing (Aigner andWoerly 2011). These treatmentsseem to result in the recovery of thefull complement of native forbs andgrasses with no need for active seed-ing. The resurgence of native forbsrepresents a tremendous food re-source for pollinators, and in thespring of 2011 we compared polli-nator activity between serpentinegrassland plots where BGG had beeneradicated and control plots that

FORBS RECOVER IN SERPENTINE GRASSLANDS WITHOUTSEEDING

were infested with BGG. Native beeabundance was more than four timeshigher in plots where BGG had beeneradicated.

CONCLUSION

These examples of grassland res-toration at the McLaughlin Reservedemonstrate that forbs play multiplevital roles in grassland ecosystems.Faced with an increasing pace ofglobal environmental change, resto-ration ecologists are realizing that itis increasingly difficult to reconstructhistorical species assemblages, andthat species for restoration shouldbe selected to meet restoration goalsunder current and future conditions.Selectively including native forbs canenhance the functionality of restoredgrasslands and make these grass-lands more resistant to invasion andresilient to environmental change.Just as importantly, including forbscan add a small reminder of themythic beauty of California’s vastprehistoric inland grasslands.

REFERENCES

Aigner, P.A., and R.J. Woerly. 2011.Herbicides and mowing to control

barb goatgrass (Aegilops triuncialis)and restore native plants in serpen-tine grasslands. Invasive Plant Scienceand Management 4: 448-457.

Heady, H.F. 1977. Valley grassland. InTerrestrial Vegetation of California, ed.M.G. Barbour and J. Major. Califor-nia Native Plant Society, Sacramento,CA.

Keeler-Wolf, T., et al. 2007. Commu-nity classification and nomenclature.In California Grasslands: Ecology andManagement, ed. M.R. Stromberg,J.D. Corbin, and C.M. D’Antonio.University of California Press, Ber-keley, CA.

Minnich, R.A. 2008. California’s Fad-ing Wildflowers: Lost Legacy and Bio-logical Invasions. University of Cali-fornia Press, Berkeley, CA.

Schiffman, P.M. 2007. Pleistocene andpre-European grassland ecosystems:species composition at the time offirst European settlement. In Califor-nia Grasslands: Ecology and Manage-ment, ed. M.R. Stromberg, J.D.Corbin, and C.M. D’Antonio. Uni-versity of California Press, Berkeley,CA.

Stromberg, M.R., et al. 2007. Califor-nia grassland restoration. In Califor-nia Grasslands: Ecology and Manage-ment, ed. M.R. Stromberg, J.D.Corbin, and C.M. D’Antonio. Univer-sity of California Press, Berkeley, CA.

Weiss, S.B. 1999. Cars, cows, andcheckerspot butterflies: nitrogendeposition and management of nu-trient-poor grasslands for a threat-ened species. Conservation Biology13: 1476-1486.

Young, S.L., et al. 2009. Functionallysimilar species confer greater resis-tance to invasion: implications forgrassland restoration. RestorationEcology 17: 884-892.

Paul A. Aigner, 26775 Morgan ValleyRd., Lower Lake, CA 95457, [email protected]; Kristin B. Hulvey, School ofPlant Biology, University of Western Aus-tralia, 35 Stirling Highway, Crawley WA6009, Australia, [email protected];Catherine E. Koehler, 26775 Morgan Val-ley Rd., Lower Lake, CA 95457, [email protected]; Leslie E. Scott, 26775 Mor-gan Valley Rd., Lower Lake, CA 95457,[email protected]

At the McLaughlin Reserve, an island ofbarbed goatgrass (Aegilops triuncialis)

marked by stakes is left for research in aserpentine meadow that has been restoredthrough prescribed burning and applica-tions of grass-specific herbicide. Barbedgoatgrass seems to be one invasive weedthat is not likely to be eliminated by reseed-ing with functionally similar native species.After removal of the invasive barbed goat-grass, serpentine grasslands return to domi-nance by native forbs, including many thatare widespread in California—in this casemountain dandelion (Agoseris heterophylla)and California goldfields (Lasthenia californica). Photograph by Paul A.Aigner.

Native pollinators, such as this sweat bee(Lasioglossum titusi) collecting pollen frommountain dandelion, are four times moreabundant in restored serpentine grasslandscompared to those infested with barbedgoatgrass. Photograph by Paul A. Aigner.

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THE CNPS GRASSLAND INITIATIVE: DOCUMENTINGGRASSLAND TYPES ACROSS CALIFORNIA

by Jennifer Buck-Diaz and Julie Evens

hat makes a greatflower year? We knowthat it has to do withclimate, particularly

rainfall and temperature, and weknow that timing is everything. Insome years, wildflower abundanceabounds in California’s grasslands,with swaths of color painting theGreat Valley floor, surrounding foot-hills, and warm deserts. Then again,there are years like 2012, when adry winter delays or even inhibitsgermination of many annual plantspecies.

A primary goal of the CaliforniaNative Plant Society’s “Grassland Ini-tiative” is to gain a more completepicture of the variation of grasslandsacross the state. With better knowl-edge of the diversity and extent ofgrassland types, we can work towardscomprehensive conservation of bothrare and common annual-dominatedplant habitats in California.

The CNPS Vegetation Programuses standardized protocols to sur-vey and classify vegetation in Cali-fornia. More than 30 new herba-ceous alliances have been recentlydescribed in the second edition of AManual of California Vegetation (Saw-yer et al. 2009), including showyannual types such as white-tip clo-ver swales (Trifolium variegatum Al-liance) and popcorn flower fields(Plagiobothrys nothofulvus Alliance).However, many grassland habitatsremain unsampled, undescribed, andoverlooked.

NEW VEGETATIONASSESSMENTS

Recent funding through privatedonations, Bureau of Land Manage-ment, California Department of Fish

and Game, Natural Resources Con-servation Service, and The NatureConservancy has allowed CNPS staffto collect additional field surveysin under-sampled areas of the state.We prioritized surveys in the SanJoaquin Valley, the neighboringCarrizo Plain, and the base of theTehachapi Mountains to fill in gapsof knowledge about grassland types.

CNPS staff collected more than400 new herbaceous surveys across2010–2011, with an additional 100surveys planned for 2012. We havealso compiled approximately 1,000surveys from previous projects andthrough collaborating researchers.Across the first two years of sam-pling, we observed an amazing array

of native perennial and native an-nual vegetation types in grasslands.Perennial types include slopes dense-ly studded with nodding needle grass(Stipa cernua), hill pockets domi-nated by California brome (Bromuscarinatus), and vast landscapes ofcurly blue grass (Poa secunda). Na-tive annual types include alkali flatscharacterized by tar plant (Centro-madia pungens), small ephemeralstream terraces prickling with thistlesage (Salvia carduacea), and bril-liantly golden slopes of commonmonolopia (Monolopia lanceolata).

We are currently analyzing bothnew and pre-existing grassland datato develop a comprehensive classi-fication of grassland vegetation in

A milky way of popcorn flower fields (Plagiobothrys nothofulvus), sky lupine (Lupinusnanus) and owl’s clover (Castilleja exserta) on the Tejon Ranch. All photographs by JenniferBuck-Diaz except where noted.

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California. However, the classifica-tion of annual vegetation can be com-plex and requires analysis beyondjust the dominance or abundance ofone species. Some annual grasslandcommunities in our state consist ofshort-lived plants that germinate andgrow quickly with cool fall/winter

rains, then bloom and set seed be-fore the summer dryness of ourMediterranean climate. A vast seedbank in the soil offers a buffer to thesystem from year-to-year variationin precipitation.

In these annual systems, it is es-sential to pay attention to indicator

species because they show a prefer-ence for certain environmental set-tings and form particular assem-blages of plants. We look closely atthese species because they specializein a particular habitat; thus, they area better ecological indicator of thatenvironment than a habitat general-ist. Indicator species may or may notbe abundant in an area or even havethe highest relative percent cover inthe grassland. An indicator may beinconspicuous such as dwarf plan-tain (Plantago erecta) which is cor-related with specific habitat types,or it may be a large, showy plantsuch as thistle sage (Salvia carduacea)which tends to be restricted to flats,terraces, and sandy washes.

A focus on indicator speciesstrengthens the classification of an-nual types by honing in on the plantsthat are indicative of a site, and de-emphasizing generalist species thatcross many different habitats anddon’t reflect the local or regionalconditions. We use statistical analy-sis tools such as Cluster Analysisand Indicator Species Analysis todetermine rigorous groups of spe-cies (communities) and the speciesthat are unique to these groups. As aresult, we are expanding the defini-tion of grassland types to includenative forbs as important, and some-

TOP: The Temblor Range of the Carrizo Plain National Monument aflame with patches ofcommon monolopia (Monolopia lanceolata) and other forbs. • BOTTOM: A vast landscape ofgrassland habitat is protected at the base of the Tehachapi Mountains on the WindwolvesPreserve, owned and managed by The Wildlands Conservancy. New surveys from this andother sites are contributing to our knowledge of annual and perennial grassland habitats.Photograph by Rebecca Crowe.

New vegetation types recently describedby CNPS staff include stands dominatedby Great Valley phacelia (Phacelia ciliata,purple) and common monolopia (Monolo-pia lanceolata, yellow). Slope, aspect, soils,and other environmental variables contri-bute to the patterns of annual vegetation.

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times dominant, plants of the com-munity.

For example, in the vegetationtype named California Goldfields–Dwarf Plantain–Six-Weeks FescueFlower Fields (Lasthenia californica–Plantago erecta–Vulpia microstachysAlliance), either one or a combina-tion of these three indicator speciesis characteristically present. A vari-ety of other native and non-nativeforbs and grasses can co-occur withthese species and may even domi-nate the herbaceous layer. The pres-ence of one or all three of the diag-nostic species within this allianceindicates a particular habitat that isoften composed of infertile soils suchas serpentine or aged clays. Whilewe still don’t have a complete pic-ture of all the associations (varia-tions in types) that fit within this

alliance, we are improving our un-derstanding of this habitat from sur-veys and analysis over the past fewyears (Buck-Diaz and Evens 2011,Buck-Diaz et al. 2012).

One exciting result of our recentdata collection and analysis is thedefinition of 16 new herbaceous alli-ances, including 5 new annual grass-land types such as annual sunflowerfields (Helianthus annuus Alliance)and virgate tarplant flower fields(Holocarpha virgata Alliance), andperennial grassland types such aswild buckwheat patches (Eriogonum[elongatum, nudum] Alliance). Wealso describe more than 50 new as-sociations, of which greater than halfare annual types. The distinctionsbetween perennial and annual grass-land are partially defined by indica-tor species such as curly blue grass

(Poa secunda), nodding needlegrass(Stipa cernua), or purple needlegrass(S. pulchra). They are also definedthrough classification analyses whichset meaningful thresholds betweenthe percent cover of perennial ver-sus annual species. These thresholdsare based on the analysis of surveydata. For example, deer grass (Muh-lenbergia rigens) has greater than 5%absolute cover while non-nativegrasses and forbs intermix with vari-able cover.

SEASONAL AND ANNUALCOMPARISONS

When classifying annual vegeta-tion types, it is also important to usecharacteristic species that have tem-poral persistence, i.e., a measure of

Photographs taken at the same site can augment repeat surveys and allow a visual documentation of annual variation in grasslands. Theleft photo column shows stands from 2010 and the right column from 2011. The top row is a flat, open plain dominated by needlegoldfields (Lasthenia gracilis) one year, and annual grasses the next. The bottom row is a hill slope dominated by common monolopia(Monolopia lanceolata) in both years, though with less cover in 2011.

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how often a species is detected overtime. If you were to visit the sameplot every spring over many years,which species will most likely bepresent no matter what the patternof rainfall or temperature? In orderto have a robust and stable classifi-cation, it will be necessary to defineannual vegetation types using spe-cies that are persistent over time.

We are beginning to revisit anumber of plots over a three-yearperiod (2010–2012) to capture boththe seasonal and the annual varia-tion in grassland vegetation. We arefortunate that the climate has variedsignificantly over this time frame,with precipitation levels rangingfrom wet (7–10 inches in the SanJoaquin Valley) to dry (approxi-mately 3 inches). Though not alldata collection and analysis is com-plete, it is apparent that some spe-cies have a higher persistence overtime than others.

Preliminary conclusions of sea-sonal sampling show that samplingin both the spring and summer cap-tures a more complete picture of na-tive forb diversity, species composi-tion, and cover. Summer surveysalone tend to over-emphasize non-

native grasses and overlook the pres-ence of ephemeral natives. Also, inextreme drought years it may not bepossible to recognize some indica-tors of annual types due to a com-plete lack of germination withinplots.

These results highlight the im-portance of timing when samplingannual stands. If only one site visitis to be completed for an area, thenthe survey should be well-timed inthe spring to fully represent thecover, composition, and overall sig-nificance of native plants. The clas-sification of annual vegetation typeswill be refined through additionalsampling across and between yearsin an attempt to utilize persistentand characteristic species.

Despite some of the amazingphotographs in this issue of Fre-montia, there is no denying thatmany grasslands in California areheavily impacted by non-native plantspecies. A semi-natural stand is theterm used for habitats where exoticspecies make up the majority (oftengreater than 90%) of the relative per-cent cover of plants and where thespecies richness (a count of the num-ber of species) is quite low. Many

non-native grassland species are gen-eralists, meaning they thrive acrossmultiple habitats and variable con-ditions, thus semi-natural stands aredescribed differently than most na-tive vegetation alliances.

Our current objective is to estab-lish baseline knowledge about natu-ral grasslands and semi-naturalstands in California. Future actionsunder the CNPS Grassland Initiativewill involve investigating the impactsof non-native plants and rangelandpractices, such as grazing with sheepor cattle, that may be used to man-age invasive plants. We hope to col-laborate with partners to conserveand restore a representation of Cali-fornia grasslands, and we will worktowards educating landowners andthe public about the values of theseunder-appreciated habitats.

REFERENCESBuck-Diaz, J., and J.M. Evens. 2011.

Carrizo Plain National MonumentVegetation Classification and Map-ping Project. Draft report preparedfor the Bureau of Land Managment.California Native Plant Society, Sac-ramento, CA.

Buck-Diaz, J., S. Batiuk, and J.M. Evens.2012. Vegetation alliances and asso-ciations of the Great Valley Eco-region, California. California NativePlant Society, Sacramento. Final re-port to the Geographical InformationCenter, Chico State University.

Dufrene, M., and P. Legendere. 1997.Species assemblages and indicatorspecies: The need for a flexible asym-metrical approach. Ecological Mono-graphs 67(3): 345-366. Great ValleyEcoregion, California. Report pre-pared for the Geographical Infor-mation Center at California StateUniversity, Chico. California NativePlant Society, Sacramento, CA.

Sawyer, J.O., T. Keeler-Wolf, and J.M.Evens. 2009. A Manual of CaliforniaVegetation. California Native PlantSociety, Sacramento, CA.

Jennifer Buck-Diaz, 2707 K Street, Suite1, Sacramento, CA 95816, [email protected]; Julie M. Evens, 2707 K Street, Suite 1,Sacramento, CA 95816, jevens@cnps. org

Tidy-tips (Layia munzii) dominate the lower Carrizo Plain while common monolopia(Monolopia lanceolata) adorns the slopes of the Temblor Range.

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CONSERVING AND MANAGING PRAIRIE, GRASSLAND,AND VERNAL POOL LANDSCAPES THROUGH

COLLABORATIONby Pelayo Alvarez

bout 250 years ago the Span-iards arrived in Californiain an event that foreverchanged landscapes in

the state. They brought with themthings from the old world that wereimportant for their survival, includ-ing livestock and a long list of plantspecies that thrived in California’sMediterranean climate. Some ofthose non-native species provide thebasis for what is now a vibrant agri-cultural multibillion dollar industry.

Other plant species, such as yel-low star thistle, were accidentallyintroduced and became invasive,wreaking havoc in both natural andagricultural systems, reducing pro-duction, fragmenting habitats, andaltering competitive interactionsamong plant species and other im-

portant ecosystem processes. Thescale and rapidity of the vegetationconversion was unprecedented andbrought about additional manage-ment challenges to the already com-plex system that comprises Califor-nia grasslands. Today only one per-cent of remaining grasslands are na-tive. Despite that catastrophic veg-etation conversion, California grass-lands remain one of the mostbiodiverse systems in the world.

ECOLOGICAL VALUE OFGRASSLANDS

There are over 11 million acresof grasslands within and encirclingthe Central Valley and the interiorCoast Range, many of which are

privately owned and managed asrangelands for livestock production.Grasslands in California providemultiple ecosystem services—thebenefits that human societies obtainfrom nature—including wildlife habi-tat, watershed protection, open space,and climate change mitigation.

California grasslands support awide variety of wildlife species in-cluding freshwater fish, birds, in-vertebrates, and mammals (Hunt-ing 2003). There are 75 species as-sociated with California grasslands,including 10 vertebrates, 14 inver-tebrates, and 51 plants that are listedas threatened or endangered underthe Endangered Species Act (Jantzet al. 2007).

California valley grasslandsarguably provide the most important

Cattle grazing is used to control invasive species and reduce fire fuel loads at Mission Peak Regional Preserve in Fremont managed bythe East Bay Regional Parks District. Photograph courtesy of Sheila Barry, University of California Cooperative Extension.

A

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wintering habitat for raptors in NorthAmerica (Pandolfino 2006). Califor-nia Bird Species of Special Concern(Shuford and Gardali 2008) whichuse these habitats extensively includeMountain Plover, Burrowing Owl,Loggerhead Shrike, GrasshopperSparrow, and Tricolored Blackbird.

Grasslands also provide impor-

tant habitat for pollina-tors that are essential forpollinating California’sagricultural crops. Thevalue of pollinator ser-vices from wild pollina-tors to California agri-culture has been esti-mated at between $937million and $2.4 billionper year (Chaplin-Kramer et al. 2011).

Despite their eco-logical value, Californiagrasslands do not enjoythe same protection thatother areas do. In the1980s and 1990s the per-

ception among land managers andthe conservation community regard-ing grazing and other ranching ac-tivities was very negative. Land man-agers in California quickly removedgrazing from land that was histori-cally grazed in an attempt to restoreit to a natural state. After a few yearsand in the absence of disturbance,

the non-native annual grasses andother invasive plant species createdan impenetrable thatch that furthersuppressed native plants. However,a few forward thinking land manag-ers began rethinking the previouslyheld views on the role of grazingwhen they saw that, on a number ofprivate ranches, the wildflowers werestill thriving in upland areas andvernal pools.

A NEW PERSPECTIVE ONGRAZING

The concept that grazing couldbe beneficial in the management ofCalifornia grasslands was not newsto the California Native Plant Soci-ety. An article in the January 1992issue of Fremontia describes howgrazing could be used to favor na-tive bunch grasses (Edwards 1992).But it was not until years later thatresearch done in California startedto provide scientific evidence that

In serpentine soils, grazed sites (right) provided betterhabitat for the endangered Bay Checkerspot butterfly thanungrazed ones (left). Photograph courtesy of Dr. StuartWeiss, Creekside Center for Earth Observation.

Cattle grazing vernal pools at North Table Mountain Ecological Preserve, managed by the California Department of Fish and Game.Photograph by Tracy Schohr.

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properly managed grazingcould be an important toolfor increasing the conser-vation value of grasslandsand associated habitats inCalifornia.

Research conductedby Jaymee Marty, an ecol-ogist with the Nature Con-servancy, and collabora-tors suggested that cattlegrazing could be an es-sential management toolin maintaining biodiver-sity in vernal pool ecosys-tems (Marty 2005). Cattleconsume the more palat-able non-native annualgrasses that invade vernalpools, allowing nativeplants and the species thatdepend on them to thrive.In addition, by reducingevapotransporation fromthe non-native annualgrasses, cattle grazing in-creases the inundationperiod of the pools by asmuch as 50 days. This al-lows vernal pool endemicinvertebrate species tocomplete their life cycle (Pyke andMarty 2005).

Similar beneficial effects of graz-ing have been found on serpentinesites south of San Francisco Bay(Weiss 1999). These sites supportmany rare plant and wildlife spe-cies, including the endangered Baycheckerspot butterfly. Studies con-ducted by Dr. Stuart Weiss, chiefscientist at the Creekside Center forEarth Observation, showed benefi-cial effects of grazing on butterflyhabitat in south San Jose wherenon-native annual grasses wereoutcompeting the dwarf plaintain, aplant that the checkerspot butterflycaterpillar feeds on. In this ecosys-tem, cattle selectively graze the non-native grasses over the native forbs,favoring the dwarf plaintain and al-lowing the butterfly to thrive (Weiss1999).

More recent research published

in the December issue of the Journalof Wildlife Management (Germanoet al. 2011) summarizes ten years ofdata on the effects of grazing andinvasive grasses on desert vertebratesin California. The study concludedthat grazing can be used to controlexotic non-native grasses and bebeneficial to vertebrate species thatrely on open habitats, such as theblunt-nosed leopard lizards, giantkangaroo rats, short-nosed kanga-roo rats, and San Joaquin antelopesquirrels, among others.

Today grazing is recognized as acost-effective and natural tool formanaging vegetation and enhanc-ing wildlife habitat and other eco-system services. It is also a tool thatis being used by several land man-agement agencies in California thatrely on local ranchers to be able tomeet their conservation goals.

Other activities associated with

ranching are also benefi-cial from a conservationstandpoint. Ranch stockponds play an increasinglyimportant role in provid-ing habitat for amphibians.In the Bay Area, stockponds on ranchlands pro-vide up to 50% of the re-maining habitat for thethreatened California tigersalamander (United StatesFish and Wildlife Service2006).

Other ecosystem ser-vices from rangelands areincreasingly being recog-nized, including openspace, watershed protec-tion, and climate changemitigation and culturalservices. Among the cul-tural values we should rec-ognize are recreation, openspace, and the knowledgeranchers have accumu-lated over generations onhow to manage theselands.

Although views on theecological importance of

grasslands and the role of ranchingare gradually changing, ranchers stillface numerous challenges to stay inbusiness. Profit margins are ex-tremely low, making conversion toother uses (vineyards, orchards) atempting proposition to ranchersthat are land rich and cash poor.Younger generations may not be in-terested in ranching, and land pricesmake it difficult for them to starttheir own operations. Little by littlethe infrastructure that once sup-ported ranching has slowly been dis-mantled, making ranching in someareas a marginal activity.

A recent rangeland vulnerabilityassessment by the Nature Conser-vancy found that every year 20,000acres of rangeland in California areconverted to intensive agricultureand urbanization (Nature Conser-vancy, unpublished data). The lossand fragmentation of these habitats

Dark blue shows rangelands of highest conservation value and levelof threat of conversion. Source: California Rangeland ConservationCoalition, 2007.

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CALIFORNIA RANGELAND TRUST

alifornia Rangeland Trust formed following around table discussion among innovative ranch-

ers who understood the need for a land trust dedi-cated to preserving not only the land, but also thestewardship and the iconic California ranching heri-tage that the ranchers provide. Since its founding in1998, the Rangeland Trust has grown substantially,and to-date has protected in perpetuity over 250,000acres of rangeland.

Rangeland Trust works to preserve grasslandsthrough voluntary conservation easements. A con-servation easement can be purchased or donated,and is an agreement between the landowner and theland trust that the land will remain as is, undevel-oped forever. Funding is applied from state andfederal programs, and more often these days comesfrom private foundations and individuals who be-lieve that the Rangeland Trust is protecting both theheritage and the future of ranching in California.Currently, the Rangeland Trust has 120 families onits waiting list that hold nearly 500,000 combinedacres of grassland.

In California, there are 34.1 million acres ofgrassland that provide sanctuary and habitat for thestate’s diverse wildlife and plant species. Of those, 22million acres are privately owned, and all are caredfor and stewarded by California ranchers.

Through the use of rotational grazing and landmanagement practices by ranchers, rangelands pro-vide habitat and homes for 95% of the state’s endan-gered and threatened species. Grazing and grass-lands are also key factors in the spectacular wild-flowers that bloom in the spring. Many of the nativeflowers would be overtaken by invasive species with-out the use of grazing as a natural management tool.

Urban development and sprawl are major chal-lenges for California’s open spaces, and they willcontinue to put pressure on grasslands and the re-sources they provide, including clean air, clean wa-

ter, and species habitat. Between 1984 and 2006 over400,000 acres of California grasslands—the equiva-lent of more than 303,000 football fields—were lostto development. With the California population stillon the rise, development pressures will continue tobe relentless.

Conservation easements held by land trusts arean extremely viable alternative to combat the pres-sures of development. Through conservation ease-ments, ranchers can receive tax benefits and fundingto ensure the future of their ranches. Many ranchersuse conservation easements to ensure that the ranchstays in the family and remains economically viable.Estate taxes often leave heirs with significant debtand few options to pay the inheritance tax. At timesthe family is forced to subdivide and sell the ranchfor development. However, a conservation easementprovides a way to pay inheritance taxes and takesdevelopment options out of the equation, loweringboth land taxes and the estate taxes.

The hard work and long hours put forth by landtrusts in California are key in preserving the state’sopen spaces and resources. Conservation easementsprovide rangelands a preferred advantage over de-velopment options. However, with the economicdownturn and state budget cuts, funds to preserverangelands are shrinking. As funds become scarcer,land trusts like Rangeland Trust will be relying moreheavily on donations from those who believe in theirmission.

For the foreseeable future, ranchers and theRangeland Trust will be facing some tough chal-lenges. However, numerous opportunities still existto preserve grasslands and the resources they pro-vide for future generations of Californians. The factthat the Rangeland Trust has a half-million acreswaiting to be protected through conservation ease-ments is a hopeful sign.

—Sahara Saude

C

reduces their ability to provide eco-system services and erodes the socio-economic framework of rural com-munities.

The accumulation of scientificevidence on the use of grazing as amanagement tool and concerns overgrassland conversion has createdan unprecedented alliance. The USFish and Wildlife Service took the

lead in bringing together ranchers,environmentalists, and regulatoryand non-regulatory agencies, afterrepeatedly hearing from the envi-ronmental community about the lossof grassland habitats and species,and from the ranching communityabout the impact that regulationswere having on their ability to stayin business.

BIRTH OF THE RANGELANDCOALITION

A barbeque in Sunol, CA pro-vided the backdrop for the first meet-ing of a group of 12 organizationswho had the courage and vision toset aside their differences and focuson potential solutions to protectCalifornia rangelands. Among them

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were representatives from the Cali-fornia Native Plant Society. “We didnot know if we were going to abarbeque or to be barbequed,” saidCarol Witham, “but it was clear thatthere was some common ground towork with.”

The new coalition was calledthe California Rangeland Conser-vation Coalition,1 and founding or-ganizations crafted a document, theRangeland Resolution, that outlinesthe principles by which Coalitionsignatories work together. A focusmap was created (see page 51) thatprioritized area in the Central Val-ley and inner Coast Ranges whereimportant species and habitatsneeded to be protected. It included500 rare plant species in grasslands,blue oak woodlands, and vernalpool habitats.

Today the Coalition has grownto more than 100 organizations, in-cluding government agencies, re-searchers, environmental organiza-tions, and others who work togetherto protect and enhance the economicand ecological values of grasslandsin the Central Valley of California.Overall the Coalition tries to pro-mote collaboration and improvecommunications among all thepeople and organizations who havean interest in these areas.

Science was essential in break-ing the barriers between the variousCoalition partners, and that madepossible the creation of this broadumbrella organization. Science re-mains central to its ongoing work,which promotes multidisciplinaryscientific collaborations and facili-tates interactions between the re-search community, ranchers, andland managers.

Outreach efforts are focused onraising awareness of the importance

of California grasslands and the needto work with private landowners fortheir protection. The Coalition orga-nizes a yearly conference, maintainsa website (http://www.carangeland.org), and hosts several field trips,workshops, and presentations on avariety of topics such as grasslandecology and management, existingconservation programs available toranchers, and emerging marketsfor ecosystem services such as car-

bon sequestration and water qualitymarkets.

In the policy arena, the Coali-tion supports public policies thatmaintain essential infrastructure andtechnical capacities of governmentagencies, and at the same time im-prove the business and regulatoryenvironment for ranchers. For in-stance, it supports policy tools thatare important to ranchers such asthe Williamson Act—which pro-vides tax incentives to agriculturalproducers in California—and theestate tax.

It also seeks common groundamong its signatories to increasefunding for grassland conservationin California. In addition, the Coali-tion works to promote collabora-tion among several governmentagencies in order to increase aware-ness of the ecological importance ofprivately owned grasslands, coor-

dinate permits for conservationprojects on private lands, and im-prove communications with theranching community.

RECOGNIZING SHAREDVALUES

Defenders of Wildlife, a nationalenvironmental organization with astrong California presence, early on

became one of the most active mem-bers of the Coalition. “We realizedthat we needed to work with ranch-ers in order to be successful at pro-tecting the native species and habi-tats that are important to our mis-sion,” explained Kim Delfino, direc-tor of the California Office of De-fenders of Wildlife. “It’s not alwayseasy for an environmental group topartner with ranchers. Sometimeswe have mistrust and preconceivedideas, but after talking with thesefolks, I’ve come to realize that weshare a love of the landscape.”

Today Defenders of Wildlife, incollaboration with other Coalitionsignatories, conducts research onidentifying and quantifying thepublic benefits of protecting Cali-fornia grasslands. A recent surveyof ranchers in the Central Valleyshowed that ranchers are indeed in-terested in protecting ecosystem ser-

FIGURE 2. RANCHER INTEREST IN SELECTED ECOSYSTEMSERVICE RELATED ACTIVITES.

Ranchers in the Central Valley were asked to rate their interest in providing ecosystemservices (Cheatum et al. 2011). Respondents expressed an interest in “improving wildlifehabitat” and “restoring native plants.”

1 The California Rangeland ConservationCoalition is an alliance of environmentalorganizations, government agencies, and theranching community whose mission is tobring together ranchers, environmentalists,and government entities to conserve andenhance the ecological values and economicviability of California’s working rangelands.

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vices such as wildlife habitat andwater quality, and increasing nativespecies (see Figure 2), provided thatthere are incentives available. Rely-ing on the ranchers’ desire to be-come more involved in conserva-tion, Defenders is now working onthe development of payments forecosystem services programs thatwill give ranchers the opportunityto stay on the land while providingall the environmental benefits thatCalifornia grasslands produce.

Other environmental organiza-tions have also come to recognizethe importance of California grass-lands and are now working withthe ranching community. By pro-viding incentives that keep ranch-ers on the land, we will be betterable to protect grasslands and theirspecies.

California grasslands and coastalprairies produce many benefits to

Californians. The protection ofCalifornia’s prairies and grasslandsrequires complex solutions and isintegrally linked to the ability ofranchers to continue being goodstewards of the land. Fostering col-laborations that cross boundariesand engage private landowners, thisbroad coalition is working towardsthe protection of these beautifullandscapes. Constantly threatenedby the worst in human nature, Cali-fornia grasslands might very well besaved by the best of it.

REFERENCES

Chaplin-Kramer, R., K. Tuxen-Bettman,and C. Kremen. 2011. Value of wild-land habitat for supplying pollina-tion services to Californian agricul-ture. Rangelands 33(3): 33-41.

Cheatum, M., et al. 2011. A Californiarancher survey on ecosystem ser-vices. Conservation Economics

White Paper. Conservation Econom-ics and Finance Program. Defendersof Wildlife, Washington, DC.

Edwards S.W. 1992. Observations onthe prehistory and ecology of graz-ing in California. Fremontia 20: 3-11.

Germano, D. J., G.B. Rathbun, and L.R.Saslaw. 2011. Effects of grazing andinvasive grasses on desert vertebratesin California. Journal of Wildlife Man-agement 75: 1-13.

Hunting, K. 2003. Central Valley grass-land habitat. In Atlas of the Bio-diversity of California. CaliforniaDepartment Fish and Game, Sacra-mento, CA.

Jantz, P.A., et al. 2007. Regulatory pro-tection and conservation. In Cali-fornia Grasslands: Ecology and Man-agement, ed. M.R. Stromberg, J.D.Corbin, and C.M. D’Antonio. Uni-versity of California Press, Berkeley,CA.

Marty, J.T. 2005. Effects of cattle graz-ing on diversity in ephemeral wet-lands. Conservation Biology 19: 1626-1632.

Pandolfino, E. 2006. Christmas birdcounts reveal wintering bird statusand trends in California’s CentralValley. Central Valley Bird Club Bul-letin 9(4): 21-36.

Pyke, C.R., and J. Marty. 2005. Cattlegrazing mediates climate changeimpacts on ephemeral wetlands.Conservation Biology 19(5): 1619-1625.

Shuford, W.D. and T. Gardali. 2008.California Bird Species of Special Con-cern. Studies of Western Birds No. 1.Western Field Ornithologists andCalifornia Department of Fish andGame, Camarillo, CA and Sacra-mento, CA.

US Fish and Wildlife Service. 2006.Designation of critical habitat for theCalifornia red-legged frog, and spe-cial rule exemption associated withfinal listing for existing routineranching activities, final rule. http://ecos.fws.gov/docs/federal_register/fr5071.pdf.

Weiss, S. 1999. Cars, cows, and check-erspot butterflies: Nitrogen deposi-tion and management of nutrient-poor grasslands for a threatened spe-cies. Conservation Biology 13: 1476-1486.

Pelayo Alvarez, 1303 J Street, Suite270, Sacramento, CA 95814, [email protected]

In the absence of disturbance such as grazing, invasive plant species dominate vernalpools, thus reducing biodiversity. Photograph courtesy of Dr. Jaymee Marty, The NatureConservancy.

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NEW CNPS FELLOW: DIRK WALTERSby David Chipping

n the fall of 1969, Dirk Waltersarrived at the California Polytech-nic State University campus in SanLuis Obispo, having just gradu-

ated from Indiana University with adoctorate in systematic botany. Hewas replacing the ailing Dr. RobertHoover, professor of botany. Dr.Hoover, who helped found the SanLuis Obispo Chapter of CNPS in1966, of course insisted that thisIndiana flatlander join CNPS at onceand learn the local flora.

Dirk became enchanted by Cali-fornia’s vegetation and soon was avery active member of the localCNPS chapter. He first became chap-ter president in 1972, and since thenhas served on the chapter board innumerous capacities including presi-dent, vice-president, recording sec-retary, and currently as chapter his-torian. He also served on the stateCNPS Board of Directors between1999 and 2001.

From 1969 until his retirementin 2004, Dirk was a professor ofbotany in the biology department atCal Poly. He continued to teach on alimited basis until 2009. Among hisacademic publications is the text-book Vascular Plant Taxonomy,which he coauthored with Dr. DavidKeil. In blending his teaching posi-tion at Cal Poly with the interests ofCNPS, Dirk arranged for the CalPoly biology department to host ourchapter meetings on campus formany years. He also encouraged stu-dents at Cal Poly to become inter-ested in native plants and join chap-ter field trips.

Dirk’s first love was teaching,and he enjoyed taking students andCNPS members on field trips intothe back country. Always stoppingto give detailed descriptions and an-ecdotes about each plant and its eco-logical relationships, his infinite pa-tience and good humor made any

Walters field trip a “must.” He stillleads field trips for the chapter, in-cluding the annual Shell Creek ex-cursion. For many years he wouldend the spring with an overnight“President’s Trip” beyond the chap-ter’s borders. As chapter board mem-ber Mardi Niles notes:

In planning field trips, Dirkcould always be counted on tolead a trip that was professional,interesting, and a lot of fun too.His trips were well planned. Healways scouted out the territoryto be visited the week precedingthe trip, and he knew what tolook for and the best possibleplace to find it.

Keeping up with Dirk couldbe the greatest challenge at thechapter’s annual field trip meet-ing at Shell Creek. Dirk didn’twant to miss anything of bo-tanical value, even in the rain.Dirk could find that special spe-cies in a huge field of flowers.Then, before you knew it, hewould be off to find another one.If you happened to spot one ofthe unique species before he did,he would always be so apprecia-tive and marvel at the find. Therewould be a gleam in his eye anda smile on his face. He reallyloved being out there and shar-ing his expertise.

Dirk has performed botanic sur-veys on behalf of CNPS’s conserva-tion program and in that role hasinfluenced planning at Hearst Ranchand other locations, including man-agement of the West Cuesta BotanicArea in Los Padres National Forest.Dirk was a member of the workinggroup and a photographic contribu-tor to the joint CNPS-City of SanLuis Obispo 2009 publication, Wild-flowers of San Luis Obispo, Califor-nia. He is currently active in photo-

graphic documentation of CarrizoPlain flora for another potentialchapter publication project. He hasalso worked to digitize old 35mmslide collections, particularly thoseof the late Dr. Malcolm McLeod, aCNPS Fellow. Visitors to CarrizoPlain National Monument will seehis digital restoration of plant iden-tification books.

Beginning in the early 1980s,Dirk and his wife Bonnie monitoredthe endangered and now federallylisted annual plant, Nipomo lupine(Lupinus nipomensis). This involvedconducting a survey and weedingnon-native plants from its habitat.In 1989 their findings appeared inthe journal Crossoma. For over 30years, Dirk and Bonnie have alsocontributed a native plant profilefor each chapter newsletter, withDirk writing the copy and Bonnieproviding the illustrations. The plantdescriptions and plant lore makethese easy reading for all ages, whileimparting a healthy dose of science.Other CNPS chapters and organiza-

Dirk Walters talking about the localvegetation to a group of mostly zoologistsat High Mountain Condor Lookout.Photograph by Steve Schubert. The bookhe is holding is Robert F. Hoover’s TheVascular Plants of San Luis Obispo County(UC Press, 1970).

I

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tions have used his material in theirnewsletters.

The plant profiles started whenDirk produced a lab manual for hisCal Poly students to replace the im-penetrable texts of the day. Dirk andDavid Keil maintained and updatedthe manual for many years. Dirk wasalso a major editor and contributorto a 1977 chapter-generated confer-ence. Proceedings were published asCNPS Special Publication No. 3, Na-tive Plants: A Viable Option.

There is almost no activity

within the San Luis Obispo chapterthat has not benefitted from Dirk’scontributions. For example, thechapter sponsors a yearly memberslide show meeting, and Dirk hasalways been the one to collect andarrange all the photos. He has alsocreated a number of other slide pre-sentations, including one designedto promote CNPS at meetings oflocal service organizations. Dirk al-most always helps to staff the “ad-vice table” at our chapter’s annualplant sale. And at local events, such

as Earth Day, Dirk has helped toassemble, host, and disassemble thechapter’s tables and displays, liter-ally for decades!

The number and variety of hiscontributions—all connected some-how to native plant science and edu-cation—boggle the mind, and weare indeed grateful for all he hasdone and continues to do.

David Chipping, 1530 Bayview HeightsDrive, Los Osos, CA 93402, [email protected]

NEW CNPS FELLOW: DON MAYALLby Santa Clara Valley Chapter Board

on Mayall, a member of theCNPS Santa Clara ValleyChapter, has been a leaderfor over 20 years in the

areas of conservation, rare plants,and invasives. Since joining CNPSin the mid-1980s, Don has been atireless advocate for serpentine plantsand habitats, and most notably inincreasing appreciation for and help-ing to preserve valuable serpentine

areas in Coyote Ridge and EdgewoodCounty Park and Preserve.

While growing up as a child inOklahoma, Don helped with hisfamily’s victory garden, and enjoyedexploring creeks in the neighbor-hood. After his family moved toSouthern California (just east of LosAngeles) when he was 12, Don con-tinued to spend his free time out-doors. Don attended UC Berkeley,

Dirk Walters and Lauren Brown on Coreopsis Hill, Nipomo Dunes, San Luis Obispo County, 2010. The yellow-flowered shrub in thebackground is coreopsis (Leptosyne gigantea, formerly Coreopsis gigantea) in its most northern occurrence. Photograph by David Chipping.

D earning a BA in psychology and anMA in sociology and then went towork for the state of California as alabor market analyst, but still lovedto be around wildlands, spendingtime in the Sierra Nevada and PointReyes. Later, while living in the Ber-keley area, Don enjoyed going oncamping and hiking trips, passingon his love of nature to his son anddaughter. During this time he

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worked in the field of demographicsand statistics, eventually retiringfrom Ohlone College in Fremont.

With his love of gardening andwildlands, it was inevitable that Donwould find CNPS, which he did in1985 by starting to buy plants at theSanta Clara Valley Chapter’s plantsale at Foothill College, and joiningin on field trips and classes. Aroundthis time Ken Himes (a future CNPSFellow) saw Don looking at photosat a chapter wildflower show andencouraged him to immediately visitthe spot where they were taken. Thislocation—which later became SanMateo County’s Edgewood Park andNatural Preserve—was slated at thetime to be an 18-hole golf courseand recreation area.

Don became the chapter’s SantaClara County conservation chair in1992, and around the same time, methis future wife Carolyn Curtis, whowas the chapter’s San Mateo Countyconservation chair. Don and Carolynworked together with many othersto organize a grassroots movementto protect Edgewood Park. Thismovement reached out to the localcommunity, government officials,and public agencies. They developeda 30-minute video that outlined thecase for saving Edgewood and whichbecame an effective publicity tool for

the effort. The video aired on thelocal cable television channel, andwas distributed to the public and tolocal agencies. The battle forEdgewood Park was eventually wonin the summer of 1993 when the SanMateo County Board of Supervisorsfinally voted to permanently protectit from development.

Inspired by Carolyn’s dedicationand persistence and the success ofthe grassroots movement to saveEdgewood, Don embarked on an-other campaign to save a threatenedregion of rare plants at Coyote Ridgein south San Jose, and protecting itfrom becoming part of the urbansprawl in southern Santa ClaraCounty. Much of Don’s involvementwith CNPS from the 1990s to thepresent has been focused on the pro-tection, appreciation, and study ofCoyote Ridge, a 7,000-acre serpen-tine area that contains numerousrare plants and animals.

Don’s professional backgroundin social sciences and statistics in-

fluenced his drive to use scientificmethods to protect Coyote Ridge.Through vegetative mapping, rareplant surveying, and working withscientists from many different disci-plines (botanists, entomologists andclimatologists), Don’s efforts haveresulted in the development of alarge base of knowledge about theserpentine habitat at Coyote Ridge.In 2004 Don also wrote a compre-hensive and effective document thathe submitted to public agencies onthe routing of high-speed rail inSanta Clara County that would havedestroyed the rare plant habitat onCoyote Ridge.

Don’s efforts to protect this largeswath of open space emphasized theimportance of responsibly openingup portions of the site for publicaccess. His ability to educate andwin over visitors to Coyote Ridgehas been integral to the develop-ment of a diverse group of support-ers, and the formation of the CoyoteRidge Committee. Don was also a

Don Mayall at CNPS Santa Clara ValleyChapter Plant Sale, April 21, 2012. Photo-graph by Judy Fenerty.

Recording data for vegetation survey, serpentine chaparral, Coyote Ridge, 2002. Photographby Sau San.

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key organizer of the vegetation sur-vey of Coyote Ridge, completed in2004 with chapter members and thestate CNPS Vegetation Program.

Over the years Don has held animpressive variety of positions withthe Santa Clara Valley Chapter, in-cluding president, vice-president,field trip chair, invasive exoticschair, plant sale chair, and WeedManagement Area representative.He has also served as the chapter’srare plants chair for Santa ClaraCounty for over ten years. At thestate level, Don has served since2004 on the CNPS Invasive ExoticsCommittee. His participation in this

committee included regular trips toSacramento to participate in theCalifornia Invasive Weed AwarenessCoalition. Don regularly partici-pated in lobbying the state legisla-ture, as well as traveling to Wash-ington and lobbying members ofCongress as part of National Inva-sive Weed Awareness Week.

Don’s clear and elegant writinghas been a valuable tool for the chap-ter in the protection of native plants.Don’s publications include severalFremontia articles, including a bookreview in January 1998 of KashayaPomo Plants, and two conservation-related articles in the Winter 2008

PETER JOSEPH CALLIZO: 1937– 2011by Jake Ruygt

eter Joseph “Joe” Callizopassed away in Napa on De-cember 14, 2011. He was awidely known and well re-

spected botanist, naturalist, and lo-cal historian who spent much of histime studying the natural areas ofNapa County, and especially all as-pects of Pope Valley where he wasraised. An early member of the Napa

Valley Chapter of CNPS, Joe servedas chapter president from 1983–86.

Joe was an only child and grewup on an 850-acre ranch that hisfamily acquired during the GreatDepression. The family raised cattle,sheep, chickens, wine grapes, wal-nuts, peaches, prunes, hay, and har-vested some timber from the ranch.They also tried their hand at mining

on the property. This “hands on”education taught Joe about horti-culture, ranching, and many build-ing trades.

His experience working withanimals on the family ranch led himto the decision to enroll at UC Davisto study zoology. After receiving hisB.A., he attended UC Berkeley wherehe received a master’s degree, also

P

Don Mayall and Carolyn Curtis (wife) at Joshua Tree National Park on a CNPS field trip,1995. Photograph by (the late) Brenda Butner.

issue, one on serpentine endemicsand the other on Coyote Ridge. Heauthored, along with Bob Case, “TheRole of the California Native PlantSociety in Invasive Exotic Plant Man-agement,” which was included inthe Proceedings of the 2009 CNPSConservation Conference.

Don has also written dozens ofarticles for the Santa Clara ValleyChapter newsletter, and for localenvironmental organization news-letters. Prior to his retirement, Donalso authored several books on ca-reer guidance, employment, and la-bor market analysis. Don has beena tireless member of the chapter’sconservation committee, and hascommented for the chapter on nu-merous CEQA documents. He wasa key member of a chapter groupthat worked on the 1997 Dow Drivelawsuit to prevent an unmitigatedloss of Santa Clara Valley dudleya(Dudleya abramsii ssp. setchellii)which was settled with the City ofSan Jose (Fremontia 36/1: 13).

Don has been an essential andinvaluable mentor to chapter mem-bers, providing guidance, scienti-fic knowledge, and institutionalmemory for the collective benefit ofboth the chapter and CNPS in itsentirety.

Santa Clara Valley Chapter of CNPS,3921 East Bayshore Road, Palo Alto, CA94303, [email protected]

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in zoology. For a time he workedthere as the head teaching assistantin the zoology department. Eventu-ally he moved to St. Helena to assisthis aging mother. He worked in thearea until acquiring a job on a vine-yard in Pope Valley, where heworked to maintain the roses plantedin the vineyards along with otherhorticultural duties. In 1987 Joe washired as the caretaker for the NapaCounty Land Trust’s Wantrup Wild-life Sanctuary in Pope Valley, wherehe lived on site for 20 years until hisretirement at the age of 70.

Joe’s involvement in local botanyspanned a period of more than 35years. In 1976 June Foote, Napa Val-ley Chapter president, contacted Dr.Gilbert Muth, a biology teacher atPacific Union College (PUC) inAngwin to establish a record of na-tive and naturalized plants of NapaCounty. Joe Callizo, Dr. Muth, andDr. Don Hemphill, professor of bi-ology at PUC developed a computerdatabase to record the distributionof native and naturalized plants inNapa County. The first edition in-cluded 618 species, but this grew toabout 1,700 entries by 1985. Joecollected hundreds of voucher speci-mens between 1975 to 1985, andcompiled thousands of geographiclocations of Napa County plant spe-cies, along with habitat, elevation,

and vegetation community data.In later years he worked closely

with Gilbert to update the computerdatabase into the first online flora ofNapa County, and assembled a fileof rare plant slides taken by numer-ous photographers in Napa County.He also helped to locate and cut thefirst trails at Skyline WildernessPark, as well as helped to start a

Native Americangarden at Bothe-Napa Valley StatePark.

In the 1980s Joeturned his attentionto the land conser-vation activities ofthe fledgling NapaCounty Land Trust,serving on its boardof trustees from1987–1997. Hestrongly supportedprotection of publicaccess to the Aetna/Oat Hill Mine Road,a popular hikingcorridor that was

improved and opened to the publicin 2009.

As caretaker at Wantrup Wild-life Sanctuary, Joe graciously assistednumerous graduate and doctoral stu-dents with research conducted there.He was also involved with the NapaValley Natural History Associationat Bothe Napa Valley State Park,hosted geologic exploration of ser-pentine habitats in Napa County,and collaborated on the writing of ahistory of Pope Valley families. Hewas knowledgeable about the localmushroom flora, and for a numberof years led walks and organizedannual mushroom fairs. His rich ex-perience living in, observing, andstudying Pope Valley for 37 yearsled to a 20-page interview by cul-tural anthropologist FeliciaShinnamon in 2007 that was pub-lished in 2011.

Joe Callizo was a gifted CNPShike leader due to his understand-ing of principles of nature, his skillin identifying species, and his senseof humor. He captivated children byhis teaching style, and impressed

Joe Callizo’s work as a botanist for over 35 years is largelyresponsible, together with faculty at Pacific Union College,for today’s comprehensive online flora of Napa County’s nativeplants.

Joe Callizo (front right) participating in a lichen keying session in 1996 sponsored by theLichen Society at the Wantrop Wildlife Preserve.

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BOOK REVIEWS

specific locations. For example, theCoast and Coast Ranges section in-cludes a description of Bear Valley inColusa county, complete with a fea-tured flower, in this case, the adobelily (Fritillaria pluriflora), driving andwalking directions, the best time tovisit, lists of flowers especially likelyto appear in colorful masses, plus otherless-common flowers that one mightalso see. Blackwell details 67 floralhotspots in all (pinpointed on a map

aspiring botanists such as myself.Joe continually developed pictorialidentification keys to the plants ofNapa County. His hundreds ofvoucher collections are preserved atPacific Union College. Joe also de-veloped the Annual Plant Watch Pro-gram that for a number of years wasutilized by many CNPS chaptersstatewide to monitor rare plantpopulations. Joe’s many contribu-tions to the knowledge of NapaCounty flora touched a great manypeople, and he will long be remem-bered.

Jake Ruygt, 3549 Willis Drive, Napa, CA94558, [email protected]

Joe standing near a large Sonoma ceanothus (Ceanothus sonomensis), a California endemic,on Hogback Mountain, March 1984.

too), providing ideas for new adven-tures for all except the best-travelledflower fans.

The bulk of this book is a month-by-month catalog of blooming flow-ers, covering most of the year. Eachchapter briefly describes what bloomsby geographical location, and provideshighlights of flowers for various re-gions. For instance, for April, the rareMount Diablo globe lily (Calochortuspulchellus) is mentioned for Mt. Diabloin the Coast Ranges. What follows is aspecies-by-species account of whatcould be expected to flower in eachmonth. Each species is listed by com-mon and scientific name, botanicalfamily, along with a brief descriptionand a photograph. Notes, habitat, and“where and when” are provided too.

For June’s swamp onion (Alliumvalidum), for example, blooming sitesinclude Sagehen Creek meadow, theWarner Mountains, King’s Canyon-Se-quoia, Mt. Eddy, Onion Valley, and theTrinity Alps. Each site includes a rangeof months when flowers could be ex-pected. Names and families correspondto the second edition of The JepsonManual. The photos for each species,all taken by Blackwell, are sharp, pleas-ing, and useful in identification.

In all, 600 species are describedand pictured. Most are herbaceous, butthere are also a number of shrubs and

Wildflowers of California: AMonth-By-Month Guide by LairdBlackwell. 2012. University of Califor-nia Press, Berkeley, CA. 588 pages.659 illustrations. $70, hardcover;$29.95, paperback. ISBN #978-0520272064.

Dr. Laird Blackwell, author of nu-merous books about flowers in vari-ous regions of California, has now writ-ten a book that will be enjoyably usedby lovers of California native flowers.While most guides to flowers are ar-ranged by color or botanical family,Blackwell’s book succeeds in a newapproach: flowers are arranged by themonths in which they usually bloom.Broader in geographical coverage thanhis previous books, Blackwell’s booklists locations especially rich in flow-ers, provides details about which spe-cies can be expected to flower in agiven month, and includes placeswhere specific flowers can be expectedto bloom.

Blackwell divides the state into tengeographical regions from the north-west mountains to the southerndeserts, and then describes easily ac-cessible sites within each area and theflowers that reliably bloom there. Eachsection opens with a geographical de-scription, plus tips on where and whento visit, followed by descriptions of

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a few trees. The author based his choiceof species on those flowers showyenough that he would go out of hisway to view them, while ensuring thatall areas of the state were represented.Non-natives are pictured too, in “bou-quets” located between the monthlychapters. Interesting and useful mate-rial is placed at the end of the book:lists of California endemics and CNPS-listed rare and endangered species areappended, and all locations mentionedin the text are cataloged and locatedon a map. References to local florasand identification guides also are in-cluded. The book closes with indicesof both common and scientific names.

There is a lot to like in this book,and it should find a well-used placein the library (or knapsack) of flowerlovers. In addition to presenting Cal-ifornia’s floral diversity in a novel,appealing, comprehensive, organized,and user-friendly way, Blackwell writessentiently and it’s clear that he loveshis subject matter. “The glorious turn-ing of the leaves eases us into winterand its joys before the flowers stir againin only a few months” is Blackwell’sgood-by to the blooming parade in thebook’s last chapter.

Not all flowers in this book will beeasy to find in their listed locations.No surprise here, as California’s florais full of endemics with specializedand limited distributions. Very rarely,some of the flowers don’t in fact occurin all of their listed locations. How-ever, this is a very minor quibble, andfinding and viewing all the plants inBlackwell’s book is certain to delightwildflower lovers.

For whom is this excellent addi-tion to California’s natural history in-tended? People who eagerly seek outflowers will doubtless enjoy using thisbook to plan trips to less familiar lo-cales. Likewise, those who desire tobegin their spring early and extend itlate will discover, as Blackwell has,that something almost always is flow-ering somewhere in our state. Anyonecurious about flowers will enjoy justbrowsing the diversity of species fea-tured in the book. Lastly, this bookmay prove useful to gardeners as asource of potential plants that wouldbroaden the seasonal appeal of theirnative plantings.

—Chris Tarp, East Bay Chapter

cause of human activities along thecoast.

The book ends with a glossary ofterms, followed by an index that in-cludes family, tribe, and scientificnames, with common names in paren-thesis. It measures 9" x 6", and is lessthan one-half-inch thick, making iteasy to carry in a backpack or tote bag,and is printed on high-quality glossypaper. This paperback field guideshould be quite useful for beachgoersinterested in the plants they may en-counter, so they may want to keep acopy in their beach bag.

—David L. Magney

A State of Change: Forgotten Land-scapes of California, text and art byLaura Cunningham. 2010. HeydayPress, Berkeley, CA. 350 pages. $50.00,hardcover. ISBN#: 978-1-59714-136-9.

This volume is a beautiful and in-formative book, as well as an inspiringread. However, my first impression ofit wasn’t positive, as I am not excitedby the common hard cover style beingcurrently used. There must be a costadvantage, with the art printed directlyon the binding as opposed to appear-ing on a removable book cover. Butwhen I started to read and examinethe artwork, I was increasingly im-pressed by the author’s many talents.She has done her homework, her art-work is impeccable, she has worked as

Beach Wildflowers of the CaliforniaTransitional Coast by Joseph Cahill,PhD. 2011. Ventura Botanical Gardens,Inc., Ventura, CA. 132 pages. $20, softcover. ISBN #978-0615449142.

The introduction to this book statesthat it is a field guide to beach plants ofsouthern Santa Barbara and VenturaCounties. Four general plant commu-nities found along the coast are de-scribed: Coastal Strand, Coastal SaltMarsh, Coastal Sage Scrub, and Ripar-ian Wetland. A map of the part of thecoast covered by the guide is provided,with place names.

The guide contains brief descrip-tions of 146 native and nonnative spe-cies. These are arranged by plant fam-ily, sort of phylogenetically, in otherwords they are grouped such that thoseclosely related are placed near one an-other. However, while the conifer fam-ily is listed first, the guide then jumpsto the monocot families instead of thedicots, as is traditional. Regardless, asa scientist, I prefer phylogenetic ar-rangements so the reader is remindedof the relationships of plants to oneanother. Most field guides are arrangedalphabetically or by flower color orgrowth form (or some combination ofthese).

Each plant description includes ascientific and common name, life form,size, and the habitat in which the plantgrows, and these are enhanced by bothuseful and beautiful photographs and/or color scans. However, the scientificnames do not seem to follow any oneparticular reference, such as the 1993Jepson Manual of Higher Plants of Cali-fornia or the more up-to-date Flora ofNorth America, but rather come frommultiple sources, some outdated. Thebook also contains a few misspellingsof plant names and factual errors. Forinstance, California buckwheat (Erio-gonum fasciculatum var. fasciculatum)is discussed; however, it is not nativeto the region (var. foliolosum is thecommon one found here).

This field guide covers 87 nativeand 59 nonnative species, which is amuch higher percentage of nonnativesthan is found in California or justVentura County (both with only about25% nonnatives). This is in part a re-sult of the invasion of this coastal habi-tat by nonnative plants, mostly be-

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LETTERS TO THE EDITOR

SAN BRUNO MOUNTAINHCP: FLAWED PLAN

Patrick Kobernus’ article on theSan Bruno Mountain Habitat Conser-vation Plan (HCP) in the previous is-sue of Fremontia (38/4, 39/1: 10–17)dismayed members of the Plan’s Tech-nical Advisory Committee (TAC). Anarticle in the October 1993 Fremontia(21/4: 11–14) by a TAC member (Sigg)can serve as a 19-year-in-advance re-buttal, and is more detailed than thisletter. The thrust of the 1993 articlewas that the fate of the then two (nowthree) listed butterflies depends on thehealth of the mountain’s ecosystem,which was declining.

The advice was ignored, and theecosystem is still declining—onlyfaster. Now that the TAC has con-vinced San Mateo County (Plan Op-erator) and the HCP Trustees that acourse change is mandatory to save

the listed species, there is a small rayof hope that effective action can stillbe taken to preserve the mountain’secosystem, its natural processes, andthe existing species.

The HCP was supposed to restorehabitat equal to that destroyed by de-velopment, but it was badly flawedfrom the start. Development began be-fore any successful restoration couldbe demonstrated. The plan lacked per-formance standards and a solid scien-tific basis, and it was—and is—grosslyunderfunded. Further, in a conflict ofinterest that would span decades, Tho-mas Reid Associates (TRA), the con-sultant that prepared the environmen-tal review of the HCP, secured onecontract after another as Plan Managerbecause their work went unreviewed,and no other bidder could compete.

As a result, we have lost hundredsof acres to development since the HCPbegan, but not a single acre has been

restored. Indeed, the Kobernus articlecondemns itself:

Management of invasive species toprotect the endangered species habitaton the mountain has been largely suc-cessful over the past 30 years. However,coastal scrub succession, in combina-tion with expanding populations of in-vasive species, continue to overtakegrassland habitat on the mountain.

Kobernus portrays the mere pres-ence of the listed species as success.TRA’s self-serving annual reports didthe same for decades before any peerreview was conducted. That peer re-view was scathing (Longcore 2004).

A major reason for the robust ex-pansion of invasive species is that TRAstaff lacked requisite knowledge of theyoung and evolving field of ecologicalrestoration. Staff was fixated on large,obvious targets such as gorse andbroom. Precious funds were spent tospray, burn, chain, or bulldoze—but

a professional biologist and traveledthroughout California, and she writesof her observations and experienceswith considerable intelligence, thought-fulness, and grace.

The book is extremely well illus-trated, with at least 75 different colorillustrations in watercolor, oil, colorpencil, or pastel. There are also fieldsketches and other simpler illustra-tions throughout in pencil, watercolor,or ink.

The book’s subtitle, ForgottenLandscapes of California, is aptly cho-sen. The author considers differentlandscapes in the state such as grass-lands, marshes, rivers, and forests, andresearches what they would have beenlike in past ages. For each she paints apicture in words, and literally with avariety of artistic media. As a botanicalartist I am impressed by the quality ofher line work, shown especially onpages 116–117 in a plate of drawingsof different grasses found in California’sgrasslands. Her work on mammals,fish, and landscapes appears to beequally precise.

My favorite renderings are thosewhere she shows a present day photo-graph—for example of the El Cerrito

Plaza in the East Bay region, or the SanFernando Valley in Southern Califor-nia—and juxtaposes it with her paint-ing of what it might have lookedlike “pre-Caucasion Invasion.” I thinkLaura Cunningham isn’t alone in imag-ining these scenes. Don’t many of usdo just the same?

The negatives of this book are few,and mostly have to do with editorialchoices. There are a few illustrationsscanned from drawings on lined paperthat I would have had redrawn or ed-

ited out. The design of the book is veryattractive, but the text for the mostpart extends across the entirety of thepage, making the reading slower. Herwriting is so substantive yet fun toread that I don’t want to be sloweddown by design!

She uses common plant and ani-mal names throughout and relies uponan appendix for scientific names. Whileit appears to be thorough, I wouldhave preferred if at least some of thescientific names appeared in the textor along with the illustrations, at leastparenthetically. It should also be notedthat some landscapes aren’t consid-ered, in particular those of mountainsand deserts.

Native Plant Society membersshould be aware that the author isinclusive in her treatment of biota: atleast half the book pertains to faunaversus flora, but this is as it should be.Her thoughtful consideration of thoselandscapes that have changed most inthe beautiful state of California willserve to educate those who are notyet familiar with the state, and inspirethose who know this state’s amazinglandscapes well to take another look.

—Linda Ann Vorobik

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with no follow-up and poor, if any,data collection. Expensive and experi-mental growing plots were created andabandoned, again without data.

Amid this confused floundering,and unacknowledged by the Plan Op-erator, newly introduced invasiveswere converting native perennial grass-lands to weedy forbs or shrubs. TRA’sreliance on five-year plans to achievespecific results didn’t allow for adapt-ing to rapidly changing conditions.When the TAC finally took over forTRA to plan the operating budget andwork scope, the planning shifted to anannual basis, allowing for immediatereaction to ever-changing conditions.

Yellow oxalis (Oxalis pes-caprae)is considered the number one threat,due largely to a hard-to-kill bulb andrapid dispersal that can convert intactareas into oxalis monocultures. TheTAC recognized its destructive capac-ity and recommended devoting an en-tire year’s discretionary budget to it,while TRA did not recognize the prob-lem. It now infests hundreds of acres.

This letter mostly addresses biol-ogy, but the HCP’s dismal results in-volve wider policy issues. Those arethe subject of an international report(Bean, Fitzgerald, and O’Connell1991). Put policy and biology together,and the San Bruno Mountain HCP hasfailed to achieve its stated mission.The Kobernus article attempts to put apositive spin on the poor execution ofa flawed plan.

REFERENCESLongcore, T., C.S. Lam, and J.P. Wil-

son. 2004. Analysis of butterfly sur-

vey data and methodology from SanBruno Mountain Habitat Conserva-tion Plan (1982–2000): Status andtrends. University of Southern Cali-fornia GIS Research LaboratoryTechnical Report No. 1. Los Ange-les, CA.

Longcore, T. 2004. Analysis of butter-fly survey data and methodologyfrom San Bruno Mountain HabitatConservation Plan (1982–2000):Survey Methodology. University ofSouthern California GIS ResearchLaboratory Technical Report No. 2.Los Angeles, CA.

Bean, M.J., S.G. Fitzgerald, and M.A.O’Connell. 1991. Reconciling con-flicts under the Endangered SpeciesAct: The habitat conservation plan-ning experience. World WildlifeFund. Harper’s Graphics/St. Mary’sPress, Waldorf, MD.

Members of the San BrunoMountain HCP Technical

Advisory Committee:

Jake Sigg, Conservation ChairYerba Buena Chapter, CNPS

Doug Allshouse, PresidentFriends of San Bruno Mountain

Ken McIntire, Executive DirectorSan Bruno Mountain Watch

OVERALL AN EFFECTIVEPLAN: THE AUTHORREPLIES

While critics such as Jake Sigg andothers have made claims that the SanBruno Mountain Habitat ConservationPlan (HCP) is a failure, those that ac-

tually visit San Bruno Mountain cansee with their own eyes that this is notthe case. Through the HCP, 90% ofthe habitat of three endangered but-terfly species has been managed forover 30 years, and all three species arestill locally abundant on the moun-tain. The conservation area also in-cludes several rare plant species andnative plant communities that havebeen protected and managed success-fully through the HCP program.

The HCP’s primary focus has beenon the maintenance of the existing habi-tat and native plant communities ofthe mountain. This strategy has beeneffective overall and was implementeddue to the greater importance of pro-tecting the conserved endangered spe-cies habitat and native plant commu-nities, and due to the higher cost ofrestoration work. Unfortunately, thesuccess of this work has been over-shadowed over the years by the harshand often inaccurate criticisms by afew highly vocal people such as Sigg.Sigg has used relatively small areas(such as the restoration work on cutslopes adjacent to the developments,or the gorse infestation in the saddle)as his examples that the HCP is a fail-ure (Fremontia 21/4: 11–14). However,these areas comprise less than 5% ofthe total area of the HCP conservationarea (San Mateo County, 2008), arevery expensive to manage, and havenot been the primary focus of the habi-tat management for nearly two decades.

Over the last three years, missionblue butterflies from San Bruno Moun-tain have been used to reestablish anextirpated mission blue population atTwin Peaks in San Francisco (Schreiber

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2012). The populations of the endan-gered callippe silverspot and San Brunoelfin butterflies on San Bruno Moun-tain are similarly robust enough to beused as source populations to seedother sites in the Bay Area where thesespecies have also disappeared. Thesespecies would not be faring well andwould possibly be on the verge of ex-tirpation, were it not for the consistenthabitat management work funded andimplemented through the HCP overthe past 30 years. It should be notedthat this habitat management work hasalso protected all of the rare plant pop-ulations in the conservation area.

Yellow oxalis (Oxalis pes-caprae)is without doubt (as Sigg points out)a significant problem on San BrunoMountain, and I agree that the HCP

habitat managers need to find moreeffective control methods. However, amore pervasive threat to the endan-gered species and the native grasslandhabitat is the process of coastal scrubsuccession, which is overtaking grass-land at the rate of approximately fiveacres per year. This process is occur-ring in many areas of California andelsewhere where grazing and/or burn-ing is no longer allowed or used.

I provided documentation on theimpact of coastal scrub succession onbutterfly habitat on the mountain inthe 2008 San Bruno Mountain HabitatManagement Plan, in an article in theJournal of Insect Ecology (2010), andin Fremontia in 2012 (Fremontia 38/4,39/1: 10–17). Though the San BrunoMountain Technical Advisory Com-

mittee (TAC) has been aware of thesuccession problem for five years, theyhave not focused any serious effort onaddressing this issue.

San Bruno Mountain is actually ina unique situation in that there is fund-ing and a habitat management plan inplace to address brush succession andinvasive species infestations consis-tently over the long-term (i.e., de-cades). Sigg and the other activist mem-bers of the TAC need to focus theirefforts on addressing the biologicalthreats to the endangered species habi-tat and the ecosystem as a whole, ratherthan continually criticizing perceivedmissteps in the past that have littlerelevance to the conservation of theendangered species and the San BrunoMountain ecosystem today.

We all care deeply about conserv-ing San Bruno Mountain. Many dedi-cated people, including biologists,planners, environmentalists, and land-owners, have worked very hard fordecades to protect its ecosystem. Work-ing together we can accomplish a greatdeal more than when we do not.

Patrick KobernusWildlife Biologist

Coast Ridge EcologySan Francisco, CA

REFERENCESLongcore, T. et al. 2010. Extracting use-

ful data from imperfect monitoringschemes: Endangered butterflies atSan Bruno Mountain, San MateoCounty, California (1982–2000) andimplications for habitat management.Journal of Insect Conservation 14: 335–346. http://www.springerlink.com/conten t /650207q10251p553 /fulltext.pdf.

San Mateo County Parks Department.2008. San Bruno Mountain HabitatManagement Plan, 2007. Prepared inSupport of the San Bruno MountainHabitat Conservation Plan. Revised2008. San Mateo County Parks andRecreation Division, Redwood City,CA.

Schreiber, D. 2012. Endangered mis-sion blue butterflies return to SanFrancisco. The San Francisco Exam-iner, April 30. http://www.sfexaminer.com/local/2012/04/endan-gered-mission-blue-butterflies-re-turn-san-francisco.

Telos Rare Bulbs

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

❏ Charge my gift to ❏ Mastercard ❏ Visa Added donation of:

Card Number TOTAL ENCLOSED:

Exp. date

Signature

Phone

Email

Join Today!

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

❏ Enclosed is a matching gift form provided by my employer

❏ I would like information on planned giving

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

NAME

ADDRESS

CITY STATE ZIP

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

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

CONTRIBUTORS (continued from back cover)

CORPORATE /ORGANIZATIONAL

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

SUBMISSIONINSTRUCTIONS

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 AdvisoryBoard

Susan D’Alcamo, Jim Andre,Ellen Dean, Phyllis M. Faber,Holly Forbes, Dan Gluesenkamp,Brett Hall, Todd Keeler-Wolf,David Keil, Pam Muick, BartO’Brien, Roger Raiche, TeresaSholars, Greg Suba, Dick Turner,Mike Vasey, Carol Witham

Megan Lulow is codirector of science and stewardship for the Irvine Ranch Con-servancy, and has been with the organization since 2006.

Roger Raiche is a landscape designer, field botanist, and author with a long his-tory of cultivating native plants. Roger was named a CNPS Fellow in 2012.

Jake Ruygt is a horticulturalist and botanist in Napa County. He has served as theconservation chair for the Napa Valley Chapter of CNPS since 1982, and workedwith Joe Callizo on the online Napa Flora.

Paula M. Schiffman is a biology professor at California State University, Northridge.She conducts plant ecology research at Carrizo Plain National Monument andSanta Monica Mountains National Recreation Area.

Leslie Scott is a restoration specialist at the UC Davis Donald and SylviaMcLaughlin Reserve, working primarily in the Eticuera Creek Watershed Ecosys-tem in Napa County.

Robbin W. Thorp is an emeritus professor of entomology at the University ofCalifornia, Davis. He has conducted research on bees for more than 55 years.

Gene R. Trapp is professor emeritus at the Department of Biological Sciences,California State University, Sacramento. His focus is field biology.

Linda Ann Vorobik is a professional botanist and botanical artist (especially TheJepson Manual), and a past editor of Fremontia. News of her workshops, shows,and lectures can be found at www.VorobikBotanicalArt.com.

Truman Young has conducted plant ecological research in various countries overthe past 35 years, and has served on the faculty of the University of California,Davis since 1996.

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CONTRIBUTORS

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

Address Service Requested

Nonprofit Org.

U.S. PostagePAID

A.M.S.

FROM THE EDITOR

(continued on inside back cover)

Paul Aigner is resident codirector of the UC Davis Donaldand Sylvia McLaughlin Reserve in Napa, Lake, and YoloCounties, where he has studied and managed serpentine andnon-serpentine grasslands for the past ten years.

Pelayo Alvarez works for Defenders of Wildlife as the con-servation program director for the California RangelandConservation Coalition.

Jennifer Buck-Diaz is a vegetation ecologist with the Cali-fornia Native Plant Society where she surveys, classifies, andmaps vegetation of California.

David Chipping is president of the San Luis Obispo Chap-ter of CNPS and past conservation director for CNPS.

Ellen Dean is curator at the UC Davis Center for Plant Di-versity. She has been instructing students in plant identifica-tion and plant collecting for over 20 years.

Julie Evens is the Vegetation Program director for CNPS.

Judy Fenerty is the Santa Clara Valley Chapter delegate tothe CNPS Chapter Council.

Glen Holstein is a retired botanist specializing in landscapeecology. He is also volunteer botanist and Chapter Councilrepresentative for the CNPS Sacramento Valley Chapter.

Kristin Hulvey is currently a post doctoral researcher at theUniversity of Western Australia where she is studying wood-land restoration.

Catherine Koehler is resident codirector of the UC DavisDonald and Sylvia McLaughlin Reserve and executive direc-tor of the Lake County Land Trust.

he theme of this issue of Fremontia concerns California’sprairies and grasslands, one of numerous native plantcommunities that are but a remnant of what they once

were 25, 50, or 100 years ago. It seems as though so much ofwhat many of us do on behalf of CNPS concerns determin-ing the condition and extant of threatened native plantspecies and sensitive habitats, and attempting to protectthem from further decline.

Humans are largely responsible for this decline. Weexploit natural resources for basic human needs and forconsumables. But too few of us pay attention to the effectour actions have on the environment. Fewer still make theconnection between an eroding environment (polluted wa-ter, air, soil from toxic chemicals) and cumulative impactsto human health (cancer, birth and immune system defects)or to plants and animals (disease, acid rain, increased toxinsaccumulating in the food chain).

Nature cannot protect itself from what we humans do tothe environment, but we can. In April 2010, in the wake ofthe failed December 2009 United Nation’s (UN) Copenhagenclimate summit, more than 32,000 scientists, environmen-talists, and others from around the world assembled inBolivia for a World People’s Conference on Climate Change.The outcome of that meeting was a proclamation called“The Universal Declaration of the Rights of Mother Earth”that participants advocated be adopted by the UN.

The Declaration acknowledges the need for a new para-digm in how we humans relate to the natural world. It recog-nizes, as have many indigenous cultures, that all living thingshave a right to well-being (i.e., clean water, air, soil), and thathumans are obligated to act to preserve these rights. It is afascinating document. I encourage you to read it in its en-tirety, at http://therightsofnature.org/universal-declaration/.

—Bob Hass

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